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MADRONO

A WEST AMERICAN JOURNAL OF
BOTANY

VOLUME IV
10S e936

Published by the

California Botanical Society Inc.
4004 Life Sciences Building, University of California, Berkeley
North Queen Street and McGovern Avenue, Lancaster, Pennsylvania

Board of Editors

Hersert L. Mason, University of California, Berkeley, Chairman.

LeRoy Asrams, Stanford University, California.

Lincotn Constance, University of California, Berkeley.

Hersert F. Copetann, Sacramento Junior College, Sacramento, California.
Artuur W. Havpt, University of California at Los Angeles.

Business Manager

Davm D. Kecx, Carnegie Institution of Washington, Stanford University,
California.

CONTENTS

PAGE
Baxer, Mito S., An Undescribed Species of Viola from Utah ............ 194
Buake, S. F., Tracyina, A New Genus of Asteraceae from Northern Cali-

THON ALOWIST hace US ND ceetig NORRIE i nage A AS a oe an 73
BuiopcGett, Cuartes O., and Meuteuist, G. L., Color Variation in Delphin-

AUN EAT CIAL CWE LOO KM iiinr a0 iectoca ta fie elas leas MMW culo ak aeerngle ey arg al vai ot ae 231
ISBACEEUNG: MURS.) (le rc yones, Mext aise) es i Wise lae sleet soby fs ao sea lhealeiaiea a (ee
ButrerFrietp, H. M., The Introduction of Acacias into California ......... 177
CiLoxey, Ina W., A New Penstemon from the Charleston Mountains, Nevada 128
CLovER, Eizapa U., Vegetational piney ¢ of the Lower Rio Grande Valley,

“1 RDB GASH SS Med aren ape ita PL ena ue nO Meena a OR a EACH Oe Al, TV
ConsTANcE, Lincotn, Reviews ................. 37, 38, 39, 132, 167, 197, 237
CoreLAND, Herspert F., The Reproductive Structures of Pleuricospora .... 1
CoreLAND, Hersert F., On the Pollen of the Mimosoideae and the Identity

of the, Supposed) Alga Phytomorula (in) se Gh che She eins 120
CoreLANnD, Herszert F., The Structure of Allotropa ...................... 137
Cory, V. L., The Occurrence of Talinum pulchellum in Texas ............ 67
NE TEBE EAM ECO VICE, cite wie pets ce uN eetiria s OTL Rice. eal k abalia yA Mala tie 196
Dayton, Witt1am A., A Cranberry from the Tahoe National Forest ...... 201
Early California Place Names Used by Daniel Cleveland ................ 67
EpPiine, Cart, Notes on Stachys rigida Nutt. ..........................4. 270
TENTETEVATRN, Nt 25 SPE Scan a Sa UA hs CE 308
Fosserc, F. RaymMonp, Eriogonum Abertianum and its Varieties ......... 189
Ganv_ER, Franx F., Notes on the Flora of San Diego County, California .. 33
GanveER, Franx F., Notes on Some San Diego County Endemics ......... 163
GarRDNER, NatHanieLt Lyon, A New Species of Chaetomorpha from China 28
Gtocx, Watpo S., Observations on the Western Juniper ................. 21
Happock, Puivie G., Picea Breweriana in Shasta County ................ 176
Hartwe zt, Rouru, Battarea phalloides (Dicks.) Pers. in Santa Barbara .. 238
Hoover, Rozert F., A New Californian Species of Brodiaea ............. 130
LN DEE TOM OT WNBEMIN M | ue caktn 8) Peng cs Sa en mM, oe ONE i ou a a 291
JEPsoN, Wituis Linn, Viae Felicitatis: The Beginning Years of the Cali-

HOLM OLATILC AM SOCIELY) ent Me ics U Loh e cite’ we lachte, segs AML G save bes 276
JouHnson, ArtHur M., Seedlings from Polyembryonic Seeds of Eugenia

UO OCT gee cr cP Ute, tpt a MME cis) ive sgn, Weal ren, Glues a teeta ee HOS 115
JonEs, GEorcE NeEvitte, New Records of Vascular Plants in Washington .. 34
McMinn, H. E., Ceanothus thyrsiflorus: Extension of Range ............ 199
Mason, Hersert L., Two New Species of Linanthus from Western North

Je\JTBL SY KEE) al eT Me CUM Rael ay EATON RL SII COT ON eee NG ee LT eR 157
Mason, ineperyT Jy, Larry Stanley Yates. ..000 000.000 0.00.. vec ee Oe 187
INMESON ERERBERT: Us., )TREVIEWS! 00 i000 os ei bee ed a Ve ve als 198, 236, 237
Mason, Herzert L., The Flowering of Wolffiella lingulata (Hegelm.)

1 Wyse NTN ay rn er ON PD A TA 241
Mason, Hersert L., A Hybrid ECO FOTN GI cs i Ns SA) aa 290
Marutias, MiIbprep, "Reviews SEATS Oe HENAN a. ren Seg REN RE | oc HARON 196
Mentauist, G. L. and Biopcetr, Cuarres O., Color Variation in Delphin-

ITA CATCINALC MAO OK. Mien aticae tian nlc uarei Mise peruit! eats, Sumo es nar alate, ge 231
Mmoy, N. T., Phylogenetic Relations of Pinus Jeffreyi and Pinus ponderosa 169
Morrison, JoHN L., Studies in the Genus Streptanthus Nutt. I. Two New

Speciesiimy the Sectiony Puclisia Nartts0 da. tiie sacs we eee Se 204
MvenscuHer, W. C., Additions to Our Knowledge of the Flora of Mount

PS AKST wi WASH TOM ec ae NU W eb Tie se Ue TMG ic ai Ma te ails le Ne 263
Nietsen, Ettar L., The Identity of Amelanchier florida Lindley ......... 17
INGEESe rN NEMS A esi ein un NG Teal. 0 alas 69, 101, 134, 168, 238
PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY ......... 72, 103, 199, 239
Quick, CLARENCE R., Notes on the Genus Ribes in California ............ 286
TRUDI a enh acide a ag AG A Pe ce MO 37, 132, 167, 236

ili
Ose

ov - 2!

Ro.urns, Reep C., Glaucocarpum, A New Genus in the Cruciferae ........ 232
St. JoHn, Harotp, A New Thalictrum from Mount Rainier, Washington .. 114
SETCHELL, Witt1aAm A., Nathaniel Lyon Gardner ........................ 126

SHarsmMitTH, Cart W., On the Identity of Claytonia nevadensis Watson ... 171
SuarsmitH, Heten K., The Native Californian Species of the Genus Core-
ODSIS Plas hie os eee eS ee Fiala od Me, & ie ak evoke telco aactie hog hee a 209
Sureve, Forrest, The Vegetation of the Cape Region of Baja California .. 105
Stespins, G. Lepyarp, Jr., An Anomalous New Species of Lapsana from

(6! 51) ot a a nr renee nO Miemn Aine INN UEP Me te ie Ge Gc ao 4 154
STEEBINS, Gi) LEDYARD, WR., Reviews oslo onc) ach lok to Rie 199
Stessins, G. Lepyarp, Jr., The Western American Species of Paeonia ... 252
STOCKWELL, Parmer and Wiccins, Ira L., The Maritime Franseria of the

Pacifie (‘Coasts ic eis oe Gk wal level, Hs Sue toe ae ee nee ar 119
Wueeter, Louis C., Eremocarpus Bentham: Preoccupied? ............... 272
Wicerns, Ira L. and Stockwett, Parmer, The Maritime Franseria of the

Pacific: ‘Coast. he e0 8 i a lhe ee As eos la ee 119
Wiecins, Ira L., Hanging Gardens of the Canary Island Date Palm ...... 260
Witson, Atzert, The California Nutmeg Tree in Cultivation ............. 166

iv

“Wr

VOLUME IV NUMBER 1

MADRONO

A WEST AMERICAN JOURNAL OF
BOTANY

he,
ut
v ( 3 Ls
Se

Contents
Tue Repropuctive Structures or Preuricospora, Herbert F. Copeland .... 1
Tue IpENTITY OF AMELANCHIER FLORIDA LinpLEy, Htlar L. Nielsen ........ Ve
OBSERVATIONS ON THE WESTERN JUNIPER, Waldo S. Glock ................. 21
A New Species oF CHAETOMORPHA FROM CHINA, Nathaniel Lyon Gardner ... 28

NoTEs ON THE F' Lora oF San Dieco County, Catirornia, Frank F. Gander .. 33

New Recorps oF VascuLar PLants In WASHINGTON, George Neville Jones .. 34

Reviews: Rollins, The Genus Arabis L. in the Pacific Northwest; Gilkey,
Handbook of Northwest Flowering Plants; Henderson, The Early
Flowering Plants in Lane County, Oregon, in 1934; Jones, A Botanical
Survey of the Olympic Peninsula, Washington (Lincoln Constance) ... 37

Published at Lime and Green Streets, Lancaster,
Pennsylvania

January, 1937

ail?

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asrams, Stanford University, California.

Dr. Lincotn Constance, State College of Washington, Pullman, Washington.
Dr. H. F. Copetanp, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Dr. P. A. Munz, Pomona College, Claremont, California.

Business Manager—Dr. Davin D. Kecx
Lime and Green Sts., Lancaster, Pennsylvania
or
Carnegie Institution of Washington
Stanford University, California

Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

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Published at Lime and Green Streets, Lancaster,
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CALIFORNIA BOTANICAL SOCIETY

President: Dr. George J. Peirce, Stanford University. First Vice-Presi-
dent: Miss Alice Eastwood, California Academy of Sciences, San Francisco.
Second Vice-President: Professor Emanuel Fritz, University of California,
Berkeley. Treasurer: Dr. David D. Keck, Carnegie Institution of Wash-
ington, Stanford University. Secretary: Miss Ethel Crum, 4004 Life Sciences
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Annual membership dues of the California Botanical Society are $2.00,
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membership should be addressed to the Secretary.

1937] COPELAND: PLEURICOSPORA 1

THE REPRODUCTIVE STRUCTURES OF
PLEURICOSPORA

Hersert F.. Coperann

The genus Pleuricospora, with the single species P. fimbriolata,
was described by Gray (7) as collected by Bolander “in or near
the Mariposa Sequoia gigantea Grove” (Mariposa County, Cali-
fornia). It is a saprophytic plant of the monotropoid alliance,
not uncommon in the Sierra Nevada; but as it is less conspicuous
than its allies Sarcodes and Pterospora, it is less familiar. It
occurs also in the Coast Range of California and in Oregon and
Washington. Two species have been described in addition to the
one usually recognized. Of P. longipetala Howell, from Oregon,
I have seen no material. As to P. densa Small (16), from the
Sierra Nevada, neither the description nor a fragment of the type
in the herbarium of the University of California shows any char-
acter by which it would be distinguished in the presence of a
good range of specimens of P. fimbriolata. AIl herbarium speci-
mens of Pleuricospora seem to have been collected during the
months June to August inclusive.

I have been led to the study of pyroloid and monotropoid
plants by a number of factors, one of which is the beauty and
abundance of several species at Jonesville, in the Sierra Nevada
in Butte County, California, at an altitude of about 5000 feet.
It is by chance rather than for any definite reason that I am now
ready to describe Pleuricospora rather than one of the others. I
have seen it in the forests summer after summer, and have pre-
pared microtome sections representing a considerable range of
stages. Ordinary microtechnical methods were used; as a fixa-
tive, usually Bouin’s fluid; as stains, Heidenhain’s haematoxylin,
Delafield’s haematoxylin and safranin, safranin and light green,
acetocarmine and aniline blue. The development of pollen was
studied in plants dug up in the mountains and brought to Sacra-
mento in the valley, where they developed rapidly; Belling’s (3)
iron-acetocarmine technique was applied, and impressed me by
its usefulness. I cannot as yet elucidate some of the most inter-
esting features of the plant; I have not worked out the detailed
anatomy of stem and root; and experimental work will be neces-
sary before one can say anything of the mycorrhiza with which
the roots are clothed, of its relation to the life of the plant, or of
the germination of the seeds.

It is a pleasure to acknowledge the many obligations which I
have contracted in pursuing these studies. I am indebted to my
father, Dr. E. B. Copeland, who made the observation of pollina-
tion noted below; to my colleagues, Dr. H. J. Child, Mr. George
Kimber, and Miss Mary Cravens; and to our students, Mr, Frank

Maprono, Vol. 4, pp. 1-40, January 6, 1937.

2 MADRONO [Vol. 4

Dutra, who identified the bee, Bremus vosnesenskiu, and especially
Mr. Taiichi Asami, who prepared the slides from which several
of the drawings were made.

GENERAL STRUCTURE

The plant is found chiefly in forests of silver-tip fir (Abies
concolor Lindl. and Gord.) in ground covered by masses of decay-
ing needles. As seen, it consists of little subglobular inflores-
cences projecting two or three centimeters above the surface
(text fig. 1). They are white in bud or young flower, later turn-

Fig. 1. Pleuricospora fimbriolata Gray. Fruiting plants in situ x %%.
Photograph by F. S. van Eckhardt.

ing yellow, brown, or black. By digging, one finds that the in-
florescences stand on peduncles several centimeters long, which
arise from masses of slender roots. The roots are usually located
in the soil beneath the decaying fir needles; their branching is, in
part at least, exogenous; but the shoots arise endogenously. In
July or August, young shoots, from a millimeter or so to several
centimeters long, are to be found among the ones already in
flower (pl. I, fig. 1) ; microscopic examination of the longer ones
shows flowers with the parts already differentiated. If one marks
the location of a cluster and digs early the next June (this is not
summer, but early spring in the mountains), one finds them some-
what more highly developed; the anthers contain pollen mother
cells. The sap of the shoots is sweet, and gives a strong positive
reaction with Fehling’s solution. In late June and along through
July, when pollen is fully formed but before the ovules are ready
for fertilization, the shoots appear above ground. The flowers
open in acropetal order, but the uppermost ones often remain
closed. Maturing and mature fruits are found in late July and
in August.

The shoots are clad with scales, of which the lower may be
called “leaves” on the “peduncle,” and the upper “bracts’”’ on the
“rachis.” They are all essentially alike, ovate to lanceolate,
sharp pointed, more or less lacerate. The phyllotaxy is irregular
and variable. Sometimes one finds a spiral, but not a typical
spiral referable to the orthodox 1/2, 1/8, 2/5, etce., system; more

1937] COPELAND: PLEURICOSPORA 3

often the scales are in whorls; on most shoots the number of
members in the whorl is constant, and is either four, five, or six:
(pl. I, figs. 1, 2).

The flowers are solitary in the axils of the bracts, and are, of
course, whorled as the bracts are (pl. I, fig. 2). In bud, the
flowers are strictly sessile; but a pedicel grows as flower and
fruit develop, and may reach a length of five millimeters. The
cluster is, then, a spike or raceme; the lower pedicels show no
tendency to bear lateral flowers, as in Hypopitys, Newberrya, and
Pityopus.

The flowers (pl. I, figs. 3-14) are hypogynous and choripeta-
lous, and with rare exceptions the parts are tetramerous. All
parts are glabrous. Of sepals, one lies against the axis, that is,
in a ventral position, and one against the bract, that is, in a dorsal
position; these two are flattened; the other two, the lateral ones,
are keeled. Sepals reach a length of 5 to 10 mm. The petals,
placed alternately to the sepals, so that two of them may be re-
garded as ventral and two as dorsal, reach a slightly greater
length. Sepals and petals, like the scales, are more or less lacer-
ate. Details of the floral diagram—maiters of the overlapping
of one sepal or petal by another—vary from flower to flower. A
stamen lies opposite each perianth segment. The anthers reach
a length of three or four millimeters, and contain distinct pollen
grains, before the filament begins to develop. As the flower ap-
proaches anthesis and finally reaches it, by the elongation of
sepals, petals, and pistil, the anther grows but little; the filament
grows to its full length, longer than the anther, in the course of a
week or so, carrying the summit of the anther to about the level
of the tips of the sepals. The flask-shaped pistil, about as long
as the sepals and stamens, bears at the base a whorl of eight in-
conspicuous blunt nectaries, which, in the mature flower, project
between the bases of the filaments. The style is short, not defi-
nitely marked off from the ovary; the stigma is narrow, its upper
surface divided into four lobes. The style is traversed by an
open channel, cruciform in cross section, and leading into the
cavity of the ovary. The ovary contains (in almost all flowers)
four placentae; these are densely covered with small ovules.
The lobes of the stigma, and the placentae, are opposite the
sepals. ‘These structures mark the boundaries of the theoretical
carpels, of which there are, therefore, four, opposite the petals.

The opening of the corolla is at first merely by a separation
of the tips of the petals, affording but a narrow passage down
to the viscid stigma (pl. I, figs. 12, 13). The flower remains in
this state for two or three days. At this stage, it is found to
have a faint, but definite, orchid-like odor; and nectar can be
found about the base of the pistil. The anthers have developed
in lateral contact with each other, and are now found to form a
cylinder about the style, not projecting above the stigma. De-
hiscence is approximately contemporaneous with the opening of

4 MADRONO [Vol. 4

the corolla. Two lengthwise rips are formed in each anther, by
the contraction of a somewhat thick-walled, but not ribbed, exo-
thecium. In fresh material, and in material preserved in liquid,
the valves are not seen to gape widely; they separate to a dis-
tance about equal to the thickness of the anther, so that the
spaces between each anther and the adjacent ones become filled
with loose pollen grains. My microtome sections (pl. II, fig. 3)
are of course made from completely dehydrated mature material,
and show the valves completely straightened out.

It is evident that pollination is by insects; presumably by
minute insects, or insects with a slender proboscis at least a centi-
meter long, to reach between the anthers to the base of the fila-
ments and pick up the pollen on the way. Only one definite
observation of a visit of an insect has been made; this was by a
bee which has been identified as Bremus vosnesenskiu (Radosz-
kowski). This is the common bee of the Sierra Nevada which is
recognized by a large white patch on the back; it has been seen
visiting other flowers of the neighborhood, both white and colored,
as Carum Gairdneri, Veratrum album, and Sidalcea spicata.

After a few days, the flower opens more widely. This is
accomplished chiefly by a movement of the two dorsal or abaxial
petals; the two petals against the rachis remain erect. The
flower is, it appears, definitely but obscurely zygomorphie (pl. I,
fig. 14). The anthers are free to fall out of their compactly
cylindrical arrangement, and may project above the stigma.
Pollination is probably accomplished, under normal conditions,
before this opening occurs; the complete opening of the flower
seems merely to be the first stage of its withering.

The fruit (pl. V, figs. 1, 2) on its gradually lengthening pedi-
cel, develops among the perianth segments. These gradually
turn dry and yellow or brown, but scarcely shrivel or lose their
shape. Fully mature, the fruit is ovoid, crowned by the shrivelled
stigma, 5-10 mm. long, and conspicuously white. It is fleshy but
utterly tasteless. Nothing is known of dissemination. The fruit
seems not to attract animals; it dries up and turns black in situ.
The abundant seeds (pl. V, fig. 3) are approximately 0.35 mm.
long. They are ellipsoid in shape and of a shining chestnut-brown
color, like minute footballs; the surface shallowly pitted by the
collapse of external cell walls; the micropylar end somewhat con-
tracted and darkened, the chalazal end obtuse, not tailed.

Tissues; Vascutar ANATOMY OF THE RECEPTACLE

On all external surfaces of the shoot, excepting those of the
anthers, the stigma, and the inside of the pistil, there is an epi-
dermis with a cuticle which is minutely striate. The ground tis-
sue shows no particular peculiarities except the presence of many
cells containing some substance (tannin ?) which stains deeply
red with safranin and deeply black with osmic acid. The same

1937] COPELAND: PLEURICOSPORA 5

substance, apparently, is abundant in the cells of the epidermis
of the anthers (the exothecium) and in the outermost cell layer
of the ovules. The stigma is seen in sections of buds to be cov-
ered by an epidermis of columnar cells; as the flower matures,
they become needle-shaped and separate from one another. They
are evidently glandular and secrete a viscid material in which
pollen grains catch.

Before describing the vascular system of the flower, one must
say something of the anatomy of the stem. There is a thin
vascular cylinder, with but few lignified cells, between the cortex
and the large pith. The “nodes are unilacunar’’; in the peduncle
there are leaf traces from each gap, but no axillary bud traces;
in the rachis, a flower trace of two bundles arises from the sides
of each gap above the bract trace.

In all of the matters just mentioned, Pleuricospora agrees yvith
Newberrya and Pityopus as I have already (5, 6) described them.
A conspicuous difference is the complete absence of epidermal
hairs in Pleuricospora.

The two bundles of the flower trace come together to form a
vascular cylinder in the pedicel. In the receptacle, several
whorls of bundles depart from this cylinder, not without con-
siderable irregularity (pl. V, fig. 5). The typical picture is as
follows: (a) First a whorl of four bundles passes out to the sepals.
(b) Then, alternating with the sepal bundles, a whorl of four
passes out to the petals. (ec, d) Next, two whorls, each of four
bundles, pass out to the stamens; the bundles to the stamens
opposite the petals are immediately above the ones to the petals;
the bundles to the stamens opposite the sepals are commonly a
little higher. This lowering of the petalad stamen bundles is
found also in Pityopus, where it is associated with the pairing of
the nectaries opposite the petals; but there is no pairing of the
nectaries in Pleuricospora. (e) A whorl of four carpel-dorsal
bundles, placed above the petalad stamen bundles, ascends the
walls of the ovary. (f) Finally, a whorl of four placental
bundles ascends into the placentae. These use up the last of the
vascular tissue.

Gaps may be either present or absent in the vascular cylinder
above any particular bundle, or elsewhere; they are most usually
present above the sepal and sepalad stamen bundles, most usually
absent above the petal bundles. No carpel-lateral bundles,
which would ascend the wall of the ovary in positions alternate
with the carpel-dorsals, were mentioned above; but occasionally
a more or less complete whorl of them may be found. Some-
times more than one bundle runs up the back of a single carpel.
The supply to a placenta consists often of more bundles than one.
My figure is based on several sets of serial sections; it represents
the ideal receptacular vascular system as I understand it, rather
than the actual structure of one individual.

6 | MADRONO [Vol. 4

DEVELOPMENT OF STAMENS AND PoLLEN

I failed to follow the development of internal structure in the
stamen from August till June. In early June, the cells of wall
and tapetum and the microspore mother-cells are already distinct
(pl. II, fig. 1). The wall consists of two or three layers of thin-
walled cells, those of the outermost layer (the exothecium, con-
taining much material stainable with safranin) being very much
the largest. |

All tapeta are short-lived; but they live long enough, in vari-
ous flowering plants, to show differences in behavior. Cooper
(4) has studied the occurrence of nuclear divisions in tapetal
cells of many species, and finds three types of behavior: either
the tapetal nuclei remain undivided; or they divide once, and the
cells remain binucleate; or they divide an indefinite number of
times, and the cells become multinucleate. No Ericales were
among the plants he studied. The present species, if my eyes
have not deceived me, conforms to none of Cooper’s three types.
By the time the pollen mother nuclei are in synapsis, the tapetum
has already begun to degenerate, but still mitotic figures may be
seen in it; mitosis seems always to be followed by cell division,
and the tapetum comes to consist of uninucleate cells of varied
sizes. The tapetum shrivels away to nothing without becoming
an amoeboid mass; it is of the “Sekretionstapetum” type of
Schnarf (18).

The development of pollen mother-cells into pollen grains
takes place, in the climate of Sacramento, during a period of a
week or two, while the anthers are about 2—3.5 mm. long.

As the anther enlarges and the tapetum degenerates, the
pollen mother-cells enlarge, and walls, gradually increasing in
thickness, appear about them. They round up. The stages of
synapsis, diakinesis, heterotypic metaphase and anaphase, and
the phases of homeotypic division were observed in succession
(pl. Il, figs. 4-8). In the heterotypic anaphase it was possible
to count the chromosomes; the haploid number is twenty-six (pl.
V, fig. 4). It was not perfectly easy to be certain of the count.
I became convinced that twenty-six is the correct haploid number
when I was able to see both anaphase groups in a single cell,
from a direction between equatorial and polar, and to find the
chromosomes corresponding in position, individual by individual,
in the two groups. They seem to be all alike, ovoid in shape,
and show a tendency to arrange themselves in definite rows.

When reduction division is complete, the wall of the pollen
mother-cell having developed considerable thickness, cell divi-
sion follows by simultaneous furrowing (pl. II, figs. 7, 8); and
soon afterward one finds separate pollen grains (pl. II, fig. 9).
Exactly this process has been described in very many flowering
plants. It raises a number of questions, to which the answers
are probably known although I have not located them. The wall

1937] COPELAND: PLEURICOSPORA 7

which develops about the pollen mother-cell, the wall which
forms in the furrows between the pollen grains as fast as they
cut in, and the walls about the separate grains, all seem to be of
the same material: at least, they are all alike resistant to stain
ing. What material is this? Does the wall of the pollen mother-
cell become divided among the pollen grains, or does it dissolve,
while each pollen grain forms its own wall?

I have not seen such a stage as Oliver (10) figures for Sar-
codes, in which the original nucleus of the pollen grain has divided
and a generative cell has formed against the wall of the pollen
grain. It is quite possible that the generative cell originates in
this fashion; I have seen it only in later stages, as a fusiform
body of deeply staining cytoplasm lying free within the pollen
grain and containing a nucleus which stains more deeply than
the tube nucleus (pl. II, fig. 9). The mature pollen grain,
mounted in liquid, is essentially spherical. The wall is marked
by four lengthwise grooves (rarely five). Mounted in air, pollen
grains become approximately cubical, as the four grooves on the
sides and also the two ends are drawn in.

As was mentioned above, the final maturing of the pollen
grains goes along with the elongation of the filaments, and is fol-
lowed by the opening of the flowers and the dehiscence of the
anthers.

Newly ripe pollen grains were tested for germination in hang-
ing drops of tap water and of solutions of sucrose of concentra-
tions from one to 30 per cent. There was no germination in tap
water. Germination took place in every sugar solution tested; it
was recognizable within five hours by the appearance of a little
bump on one of the grooves of each pollen grain. It seemed
most normal and abundant, and the growth of the pollen tube
seemed most rapid, in the 20 per cent solution; here tubes 0.5
mm. long were seen after thirty hours. The more dilute solutions
seemed hypotonic with respect to the contents of the pollen
grains; they did not cause the grains to swell and burst, but
caused the ends of the pollen tubes, where the walls must be
weakest, to become swollen, sometimes to diameters as great as
the pollen grains from which they had sprung. The more con-
centrated solutions caused no obvious damage but seemed to de-
crease the percentage of germination and to retard growth. The
generative cell was seen to enter the pollen tube before its
nucleus had divided (pl. II, fig. 10). Either the tube nucleus or
the generative cell may enter the tube first.

Of the growth of the pollen tube under natural conditions, I
know little. The pollen grains are caught on the stigma by some
viscid substance secreted by the epidermal layer of needle-
shaped cells. The pollen tubes grow along the cracks between
the lobes of the stigma and down the style channel. In the
ovary I have only occasionally been able to recognize pollen
tubes, and I have not been able to see one growing from outside

8 MADRONO [Vol. 4

into an ovule. The plugs closing old pollen tubes, mentioned by
Oliver as occurring in Sarcodes, were recognized.

DEVELOPMENT OF OvVULE AND SEED

Ovules are recognizable, up to about the time of reduction
division in the anthers, as abundant hemispherical bumps on the
placentae. I have missed seeing a series of stages including the
differentiation of the archesporial cell, the “bending over’ by
which the ovule becomes anatropous, and the origin of the integu-
ment. As the pollen grains are ripening, one finds the arche-
sporial cell (which, as is usual in Sympetalae, is itself the mega-
spore mother-cell) in synapsis or diakinesis (pl. III, fig. 1); the
integument, of two layers of cells, is closing over the nucellus,
which is a single layer of cells. Subsequently one finds a T-shaped
megaspore tetrad (pl. III, fig. 2), of three minute cells and one
large one in the chalazal position. In the latter, 2-, 4-, and 8-
nucleate stages have been observed, and finally the mature embryo
sac, consisting of an egg, two synergids, an endosperm mother-
cell containing two polar nuclei, and three antipodal cells (pl.
III, fig. 4). While the embryo sac develops, the three non-func-
tional megaspores are absorbed, and so is the whole of the
nucellus. There may be some increase in the number of cell
layers of the integument; but the integument remains quite thin,
and no jacket layer of columnar cells is formed about the embryo
sac.

It has not been possible to follow the details of fertilization.
It is clear that the pollen tube enters through the micropyle and
creates a certain amount of wreckage in the micropylar end of
the embryo sac. The egg remains clearly recognizable. In a
single section I was able clearly to see one spherical sperm
nucleus uniting with the egg nucleus, while another was uniting
with the two polar nuclei (pl. III, fig. 5). These observations
are in harmony, as far as they go, with those of Shibata (14) on
Monotropa uniflora, in which he found the sperm nuclei, elongate
when first discharged from the pollen tube, to become spherical
as they reach respectively the egg nucleus and the polar nuclei,
and only then readily stainable. Shibata (15) has mentioned the
handsome strands of cytoplasm, extending from the original
fusion nucleus in the endosperm, which he observed in living
material of Monotropa. They are equally evident in microtome
sections of Pleuricospora (pl. III, fig. 6).

The endosperm develops in the manner usual in Ericales.
Before the zygote undergoes any divisions, the endosperm nu-
cleus divides twice; each nuclear division is followed by a cell
division, the walls falling at right angles to the axis of the
embryo sac, so that a row of four cells is formed (pl. IV, figs. 1,
2). The zygote grows into the shape of a narrow tube, whose
summit, in which lies the nucleus, penetrates into the second cell
of the endosperm. Of the cells of the four-celled endosperm,

1937] COPELAND: PLEURICOSPORA 9

some or all may divide by further transverse walls; the second
cell (the one surrounding the summit of the embryo) and the
fourth (the one at the chalazal end) almost always do this; the
first and third more frequently remain undivided. By these divi-
sions the endosperm is converted into a row of six to eight cells.
Longitudinal divisions also take place, converting the cells of the
row into tiers of cells. The standard number of cells in the tier
seems to be four, but this number is not at all constant; thus one
may find a tier of four between the developing embryo and the
micropylar end of the endosperm; six or eight in a section cut
through the embryo; four in sections cut farther back; and only
one at the chalazal end.

The first division of the zygote separates, by a transverse
wall, a cylindrical cell toward the micropyle from a nearly spheri-
cal one toward the chalaza (pl. IV, fig. 3). The protoplasm in
the cylindrical cell is gathered at the end which is against the
spherical cell, and in later stages it is found to have secreted a
wall cutting off a brief suspensor devoid of contents from a coni-
cal cell which has been designated as the hypophysis. The
spherical terminal cell divides by two vertical walls into a cluster
of four. Possibly division sometimes goes farther than this, but
I have not been able certainly to recognize it. As the seed ap-
proaches maturity, stainable material accumulates in the living
cells of the endosperm and embryo. It is evidently the same
material in both structures; they are distinguishable only by
position, not by staining reactions (pl. IV, fig. 4).

At the time of fertilization the embryo sac is surrounded by
an integument partly of two layers of cells, but of three or more
against some parts of the sides of the embryo and of several
layers at the ends. Even before fertilization the outermost
layer, except for a broad gap at the micropylar end, becomes
markedly different from the others by an accumulation of tannin
(?) as a hollow vescicle within each cell. During development
after fertilization, the inner cells against the sides of the endo-
sperm become flattened and finally almost—not quite completely
—disappear. A certain number of thin-walled inner cells at the
ends remain intact into the ripe seed. A group of antipodal cells
ean be detected in the chalazal end for some time. The outer-
most layer of cells becomes very thick-walled, especially on their
lateral and inner surfaces; the tannin (if tannin it be) by which
these cells are distinguished in earlier stages is perhaps a plastic
material from which these thick walls are built. No terminal
haustoria are formed on the endosperm, and no tails are formed
on the seed by the collapse of functionless inner cells.

Discussion

The most important contributions on the microscopy of the
reproductive structures of Ericales have been those of Koch (9)
on Hypopitys; Oliver (10) on Sarcodes; Stevens (17) on Epigaea;
and Samuelsson (11) and Hagerup (8) on a variety of genera,

10 MADRONO [Vol. 4

chiefly Scandinavian. These contributions have been duly sum-
marized by Schnarf (13). Pleuricospora conforms to the charac-
ters of the order in a whole range of characters, among which
may be mentioned the following: (a) flowers formed the year
before they are to open; (b) the absence of an endothecium in
the anthers; (c) the non-amoeboid tapetum in the anthers; (d)
reduction division occurring in the anthers sooner than in the
ovules; (e) “simultaneous” division of the pollen mother-cells;
(f) pollen grains with two nuclei; (g) the channeled style; (h)
ovules with a single integument and a thin nucellus which is soon
absorbed; (i) embryo sac developed in “normal” fashion; (j) a
young endosperm of four cells in a row; (k) the inner cell-layers
of the integument absorbed, for the most part, by the endosperm.

The tapetum in which the cells continue to divide for some
time and do not become multinucleate may be an ordinal char-
acter; I know of no data on this point from other Ericales.

The pollen grains which are solitary, not in tetrads, are not
typical of the order; but they are altogether typical of the
monotropoid alliance. :

I know of no previous count of the chromosomes in any plant
of the monotropoid alliance. Hagerup counted the chromosomes
in many other Ericales and concluded that the fundamental num-
ber is six; the twenty-six chromosomes of Pleuricospora may be a
set of 4n+2. This number is not in good harmony with the
chromosome number of Pyrola (n= 238); it harmonizes well with
the numbers in Rhododendron (a genus in which Sax (12) has
found the chromosome number remarkably constant) and Ledum
(n= 18) and in the Arbuteae (n= 18 or 26).

In Ericaceae the integument is usually of several layers of
cells at the time of fertilization; and the endosperm usually
comes to consist of many cells. Most of the Pyrolaceae of
Engler and Prantl (that is, the allies of Pyrola and of Monotropa)
have a very thin integument; and in Hypopitys and Pleuricospora
the endosperm is of very few cells. The nature of the endo-
sperm in other Pyrolaceae, excepting Sarcodes, is not definitely
known. In bulk both of integument and of endosperm, Sarcodes
lies between the other Pyrolaceae and the Ericaceae.

The course of formation of the endosperm in Hypopitys and
Pleuricospora shows striking identities, notably in the transverse
divisions which take place in the second and fourth cells of the
four-celled endosperm; but Pleuricospora differs from Hypopitys
—and, indeed, from almost all other known Ericales—in the fact
that the terminal cells of the developing endosperm do not take
on a different appearance from the rest, and do not eventually
collapse. Some of the details of the embryology of Sarcodes as
figured by Oliver are different from corresponding stages of
Pleuricospora and Hypopitys; Oliver has represented an embryo
developing in the micropylar chamber of the four-celled endo-
sperm, and a suspensor of two or three short, nucleated cells.

1937] COPELAND: PLEURICOSPORA 11

Hypopitys and Monotropa are closely related to a series of
genera with parietal placentation, Newberrya, Pityopus, and Mono-
tropastrum (Andres, 1, 2). Pleuricospora stands outside this
circle, distinguished by a combination of characters of no great
weight, but which nevertheless make it seem distinct: its com-
pletely glabrous character, the pattern of the anthers, its drying
brown rather than black, its carpels almost constantly four rather
than varying about eight as an average. I do not think that it is
related to Hypopitys and its closest allies either as ancestor or
descendant, but that it is a reasonably close collateral relative.

The direction of evolution by which these plants are related
to the Ericaceae remains open for discussion. It is possible to
conceive that Pleuricospora, with its simple endosperm and anthers
opening through slits, is primitive; and that Hypopitys and its
allies represent a line of evolution leading through Sarcodes and
Pterospora to the Ericaceae, among which some of the Arbuto-
ideae might be the nearest. I am much more inclined to read the
series in the opposite direction, and to regard Pleuricospora as an
end product of a line of evolution in which many of the special-
ized structures of the Ericaceae have been lost or reduced to
their minimum essentials.

Sacramento Junior College,

Sacramento, California,
June, 1936.

LivreRATURE CITED

1. Anpres, H. Uber die Pirolaceen-Gattung Monotropastrum H. Andr.
Notizblatt Bot. Gart. Berlin 12: 696-699. 1935.

Pp Pirolaceae. Symbolae Sinicae 7: 762-768. 1936.

3. Bretitinc, Joun. The use of the microscope. New York. 1930.

4, Cooper, D.C. Nuclear divisions in the tapetal cells of certain angiosperms.
Am. Jour. Bot. 20: 358-364. 1933.

5. Copenann, H. F. The structure of the flower of Newberrya. Madrojfio 2:
137-142. 1934.

6. ———————_. On the genus Pityopus. Madrofno 3: 154-168. 1935.

7. Gray, A. Characters of new plants of California and elsewhere... .
Proc. Am. Acad. 7: 327-402. 1868.

8. HaGerup, O. Morphological and cytological studies of the Bicornes.
Dansk Bot. Arkiv 6°: 1-26. 1928.

9. Kocu, L. Das Entwicklung des Samens bei Monotropa Hypopitys L.
Jahrb. wiss. Bot. 13: 202-252. 1882.

10. Ontver, F. W. On Sarcodes sanguinea Torr. Ann. Bot. 4: 303-326. 1890.

11. Samvetsson, G. Studien iiber die Entwicklungsgeschichte der Bliiten
einiger Bicornes-Typen. Svensk bot. Tidskrift 7: 97-188. 1913.

12. Sax, K. Chromosome stability in the genus Rhododendron. Am. Jour.
Bot. 17: 247-251. 1930.

13. Scunarr, K. Vergleichende Embryologie der Angiospermen. Berlin.

1931.

14, Surpata, K. Die Doppelbefruchtung bei Monotropa uniflora L. Flora 90:
61-66. 1902.

15. ———————_.__ Experimentelle Studien iiber die Entwicklung des Endo-

sperms bei Monotropa. Biolog. Centralbl. 22: 705-714. 1902.
16. Smatzt, J. K. Monotropaceae. North American Flora 29: 11-18. 1914.
17. Stevens, N. E. Dioecism in the trailing arbutus, with notes on the mor-
phology of the seed. Bull. Torr. Bot. Club 38: 531-543. 1911.

12 MADRONO

1.0 em

Pruate I. PLevuricosporA FIMBRIOLATA Gray. Fig. 1. Young shoot, with
leaves in whorls of four, X2. Fig. 2. Rachis of inflorescence, showing attach-
ment of bracts and flowers in whorls of five, X2. Figs. 3, 4, 5. Young bud,
stamen, and pistil x5. Figs. 6, 7, 8. Older bud, stamen, and pistil. Figs. 9, 10.
Bud ready to open and stamen X5. Fig. 11. Floral diagram. Figs. 12, 13.

Open flower and longitudinal section x5. Figs. 14, 15. Flower past anthesis
and stamen * 5.

1937] COPELAND: PLEURICOSPORA 13

~v0%e* 4

{) ta oS y

ax > >3N ry a. b

cQ~ 5 fg ®t, 8 4

COS fae
ae

aes. & :

ae
(ALEKS)

Pirate II. Prevuricospora FIMBRIOLATA Gray. Figs. 1, 2,3. Cross sections
of anther in successive stages X 125. Figs. 4-9. Development of pollen grain
x 400. Fig.10. Germinated pollen grain with pollen tube attached x 400.

[Vol. 4

MADRONO

14

a Gray. Development of ovule.

Pratre III. PLEeEvuRICOSPORA FIMBRIOLAT

a2
oS
= (3)
ae
AS
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a
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celled endosperm. All x 400.

1937] COPELAND: PLEURICOSPORA 15

Prats IV. Puxevuricospora rimpriotata Gray. Development of seed. Fig.
1. Two-celled endosperm. Fig. 2. Four-celled endosperm. Fig. 3. Two-celled
embryo. Fig.4. Nearly ripe seed. All Xx 400.

16 MADRONO [Vol. 4

1.0 mm
ie Bi a TE EE

Pirate V. PLEURICOSPORA FIMBRIOLATA Gray. Figs. 1, 2. Fruits x5. Fig.
3. Seed X100. Fig. 4. Heterotypic anaphase in pollen mother cell x 840. Fig. 5.
Reconstruction of the vascular system in the receptacle x 50.

1937] | NIELSEN: AMELANCHIER FLORIDA 17

THE IDENTITY OF AMELANCHIER FLORIDA LINDLEY

Ertar L. Nietsen

In connection with a current study of the genus Amelanchier
in Minnesota, the question of the identity of A. florida Lindley
has arisen. The exact status of this species and the probability
of its occurrence in the middle western states has been a vexing
problem for some time. This has led to a study of the type
specimen and of Lindley’s (2) original description and plate;
also to a critical examination of herbarium specimens from the
region of the type collection. The material upon which Ame-
lanchier florida is based was collected by David Douglas in “‘north-
west America” in 1826. The type specimen of this species is
now deposited in the Herbarium of the Botany School, Cam-
bridge, England. The following remarks concerning the type
are from A. C. Seward’s letter to the writer:

“There are two specimens on the sheet: a smaller one, against which
is written ‘N.W. Am. Douglas.’ Below the larger is added ‘Am.
Florida’ with a reference to the Botanical Register 1833, where it is
figured and is said to be found by Douglas. In the left-hand bottom
corner of the sheet is written ‘N.W. America Douglas 1826... .
The writing is Lindley’s, though I am not by any means sure that the
words written against the smaller specimen which I quoted are in his
handwriting. The specimens are described on a more recent label as
the type of Amelanchier florida Lindley, Botanical Register 1833, Pl.
1589.”

In 1833 Lindley (2) described the species as follows:

“A. florida; foliis oblongis utrinque obtusis versus apicem grossé
serratis semper nudis, bracteis stipulisque apice plumosis deciduis,
racemis strictis multifloris, staminibus calyce extus glabro brevioribus.

“Frutex erectus, glaber, ramis viridibus v. fusco-viridescentibus.
Folia oblonga, basi utrinque obtusa, nunquam pubescentia nisi ali-
quando sub vernatione, versus apicem grosse serrata. Stipulae brun-
neae, marcidae, lineares, intus ad apicem villis longis plumosae; citis-
sime deciduae. Racemi terminales, cylindracei, multiflori, glaberrimi,
stricti; bracteis linearibus, acutis, apice villosis, citissime deciduis.
Calyx extus glaber, intus pubescens; dentibus acuminatis, staminibus
longioribus v. aequalibus. Petala lineari-spatulata. Stamina erecta,
brevissima.

“A native of North-west America, where it was found by Mr.
Douglas. It forms a handsome hardy shrub, in the way of the Snowy
Mespilus, flowering in May. Like that species, it is best propagated
by layers.

“Tt is at once recognised by the shortness of the stamens; otherwise
it is very near A. sanguinea, already figured at fol. 1171 of the pres-
ent work. But that species has moreover smaller and more capitate
flowers, the teeth of the leaves are finer, the bracteae and the stipulae
much more pubescent, and the calyx far more downy inside.

18 MADRONO [Vol. 4

“The petals vary in length; in one of our wild specimens from Mr.
Douglas they are more than 34ths of an inch long.”

Through the courtesy of the Botany School Herbarium, it was
possible for Miss Sarah Dyal, Cornell University, to examine for
me the type specimen and note certain critical features. These
can best be described in Miss Dyal’s own words (in lit.) :

“Type of Amelanchier florida Lindley 1833 collected by Douglas
1826 in NW America. (Under Aronia). Leaves glabrous; twigs dark
reddish-brown; inflorescence (a) length 5—7.5 cm., (b) glabrous, (c)
lower pedicel 8-10 mm. to base of ovary, (d) ovary summit woolly,
(e) width of hypanthium 4—4.5 mm., (f) length of sepals 3-4 mm.
(2-3 times as long as wide) woolly above, both extended and recurved,
(g) length of petals 8-11 mm.

From Lindley’s description and Miss Dyal’s observation there
seems little doubt that Amelanchier florida is an entirely different
entity from any of the several densely pubescent forms of the
Great Plains region which have been referred to that species in
the past, one of which is unquestionably A. alnifolia Nuttall (3)
often cited as a synonym of A. florida.

With this concept in mind an examination has been made of a
number of herbarium specimens from the general region of the
type which were kindly placed at the writer’s disposal by Profes-
sor M. E. Peck of Williamette University, Oregon, and Dr. T. C.
Frye of the University of Washington. It has been found that
A. florida is a well defined species occurring typically on the west
side of the Cascade Range in Washington and Oregon. All of
the typical specimens examined have a sparse evanescent pubes-
cence. This is particularly true of plants that have not yet come
into full anthesis while in the fruiting condition they are either
glabrous or with a few hairs along the midrib near the leaf base.
This agrees with Lindley’s description since the statement “‘sem-
per nudis”’ of the first paragraph is qualified in the next by “nun-
quam pubescentia nisi aliquando sub vernatione.” From the
study it has been possible to formulate the following more com-
plete description.

AMELANCHIER FLORIDA Lindley

Leaves oblong (seldom oblong-elliptic), obtuse at both ends,
fully expanded at anthesis; sparsely pubescent when young, the
pubescence quickly evanescent, early glabrate or with a few hairs
persisting on the petiole and lower midrib; blades about 4 cm.
long, 2.5 cm. wide at flowering time, about 5.5 cm. long, 3 cm.
wide at maturity, coarsely serrate toward the apex with acute
sinuses, lateral veins 10-12 (9-13) on each side ascending and
running into the serrations; petioles slender, often 1—-1.5 cm. long
at flowering time, 1—-2.5 cm. long when fully mature; inflores-

1937] NIELSEN: AMELANCHIER FLORIDA 19

re]

i
NY

Mss Drane deb, bly J Mdgwray T6 9 Feicadtilly Seire, 1, 1EF9. Y Wak. n

_ Pratt VI. Ametancuier Fioripa Lindley. An exact outline drawing of
Lindley’s original figure prepared as recommended by Buchholz (1).

20 MADRONO [Vol. 4

cence 3—7.5 cm. long, erect, at first sparingly pubescent but soon
glabrate; lowermost pedicel 8-11 (14) mm. long; sepals 3-4 mm.
long (2-8 times as long as wide), very acute, woolly above in
flower, somewhat reflexed and glabrate in fruit; petals 8-13 (16)
mm. long, linear-spatulate to oblanceolate; stamens shorter than
the calyx lobes; ovary summit woolly at anthesis, sometimes
nearly glabrate in fruit; hypanthium 4—4.5 mm. in diameter, shal-
lowly cup-shaped, only slightly constricted in fruit; fruit globose.

A. slender shrub about 15 feet high, of the forest, and more
often the forest margin, on the west slope of the Cascade Range
in Washington and Oregon.

Wiegand (4) in his first paper on the genus Amelanchier in
eastern North America reported A. florida from Isle Royale and
Keewenaw Point, Michigan. He (5) later became doubtful as to
the occurrence of this species in the Great Lakes region as indi-
cated in his second paper (5) dealing with this genus. In dis-
cussing A. huronensis he states “this species together with A.
humilis probably forms the basis of records of A. florida Lindley
from the region of the Great Lakes.” Since a number of west-
ern species of plants actually do occur about Lake Superior, a
careful study of the Isle Royale material, including some of the
specimens cited by Wiegand, was made to determine whether
Amelanchier florida has a similar distribution. A comparison of
the Isle Royale material with specimens of typical A. florida
shows the following differences: in Amelanchier florida the leaves
are oblong, coarsely serrate toward the apex, the sepals 3-4 mm.
long, 2-3 times as long as wide, very acute, the hypanthium shal-
lowly cup-shaped. In the Isle Royale material the leaves are
broadly oval to oblong-oval, usually serrate-dentate to the
middle, the sepals 1.5-2.5 mm. long, about as long as wide,
broadly acute, the hypanthium saucer-shaped. Other reports of
A. florida from the region of the Great Lakes should be dis-
counted because they apparently are based upon the earlier inter-
pretation of Wiegand (4).

Amelanchier florida Lindley appears, therefore, to be a species
of the west slope of the Cascade Range in Oregon and Washing-
ton. It should not be confused with forms of the Great Lakes or
the Great Plains regions. The following specimens, which have
been examined, may be considered as fairly typical. The letters
(UW), University of Washington, and (W), Williamette Univer-
sity, indicate the herbaria where these specimens are deposited.

Wasunineton. King County: Seattle, E. S. Meassy (UW).
Pierce County: sandy soil, Reflection Lakes, Mt. Rainier, July
31, 19382, F. A. Warren 1786 (UW); rocky soil, Gobbler’s Knob,
Mt. Rainier, July 17, 19382, F. A. Warren 1604 (UW). Whidby
Island: rocky shore, Cranberry Lake, June 2, 1934, G. N. Jones
4897 (UW) (typical except for the sepals). Orrcon. Lane
County: Horse Pasture Mt., 10 miles south of McKenzie Bridge,

1937] GLOCK: WESTERN JUNIPER 21

alt. 5000 ft., July 1, 1934, M. E. Peck 2593 (W); bank of Frog
Lake, July 27, 1927, M. E. Peck 15910 (W).

I wish to express my thanks to Dr. C. O. Rosendahl and Dr.
F. K. Butters of the University of Minnesota for their counsel
and suggestions given during the course of this investigation.

College of Agriculture, University of Arkansas,
Fayetteville, Arkansas,
March 11, 1936.

LITERATURE CITED

Bucuunouz, Arzert. Das Photo Allerlei. 32-33. Sept. 1933.
Linptey, Joun. Botanical Register, n. ser. 6: pl. 1585. 1833.
Nourratt, Tuomas. The genera of North American Plants and a catalogue
of the species to the year 1817. 1818.
Wiecanp, K. M. The genus Amelanchier in eastern North America. Rho-
dora 14: 117-161. 1912.
Additional notes on Amelanchier. Rhodora 22: 146-151.

Chics sheng yo pe

1920.

OBSERVATIONS ON THE WESTERN JUNIPER
Watpo S. GirocKx

A brief study of the western juniper (Juniperus occidentalis
Hook.) in the high Sierra Nevada of Alpine and Tuolumne coun-
ties, California, has yielded some interesting information and has
raised some important problems. The species was observed and
core samples taken in Faith and Charity valleys southeast of
Lake Tahoe, in the vicinity of Kit Carson Pass, and in the Sonora
Pass and Dardanelles region. The writer’s attention was di-
rected to the juniper by Mr. Clarence K. Bennett and Dr. Ralph
W. Chaney. Especial gratitude is due Mr. Bennett whose finan-
cial aid helped to make the field work possible.

Specific localities inhabited by the junipers are quite dis-
tinctive. These trees occupy relatively dry sunny slopes with
westerly or southerly exposure. However, scattered trees were
seen on a precipitous easterly slope whose individual jutting
ledges of nearly bare granite provided sites which have approxi-
mately the same exposure time to the sun as have the westerly
slopes. The junipers are most frequently found on gravel mo-
raines, on slopes of coarsely weathered lava, or on bare granite
ridges where their roots insinuate themselves with difficulty
among the huge boulders and into crevices. Such habitats,
rather than rich humus or moist meadow borders, are preferred.

In Faith Valley at an elevation just below 8000 feet, the trees
grow upon a weathered lava slope and also upon the side of a
gravel ridge. At Chipmunk Flat along Deadman Creek at an
elevation of 8000 feet, they grow on a coarse boulder terrace
fifteen to twenty-five feet above stream level. The north wall of
Deadman Canyon, above Chipmunk Flat, rises to a bare granite
shoulder and ridge at about 9000 feet. Here, scattered junipers,

22 MADRONO ! [Vol. 4

low, squat, and grotesquely gnarled, maintain what appears to be
a precarious foothold in the crevices. In the vicinity of lower
Relief Creek at 6500 to 7000 feet and on the upland west of the
mouth of Deadman Creek (Dardanelles Quadrangle, United
States Geological Survey) at 8500 feet the junipers grow on rock
ledges and on gravel knolls and ridges. Here, on what is called
East Flange Ridge, they probably have their best development.

On the whole the western juniper, in contrast with its rela-
tives, inhabits the subalpine zone at comparatively high eleva-
tions near timberline. The California juniper (Juniperus cali-
fornica Carr.) and the desert juniper (Juniperus utahensis Lem-
mon) are commonly among the first trees encountered above the
Lower Sonoran zone, whereas the western juniper is commonly
among the last encountered at the upper tree limit. In habit, in
form, and in age the two types differ remarkably. The high alti-
tude species possesses a more highly developed arborescent form
than its lowland kin; it is in certain cases regal in its proportions.
Its vitality is such that although the stress of weather and the
presence of rock fragments may distort and malform it into
weirdly grotesque caricatures of trees, it survives the wrack and
stress until the last bit of cambium dies.

Western junipers, on the whole, are not only solitary as a
species but also solitary as individuals. They are intolerant of
shade and of crowding. Where grouped, the trees are spaced
openly much in the manner of the western yellow pine (Pinus
ponderosa Dougl.). Many of the veterans stand completely iso-
lated on a sunny slope or knoll. Close association with other
species of trées is rather uncommon in the localities of this study.
However, in the same general area red fir (Abies magnifica Murr.)
and lodgepole pine (Pinus Murrayana Balf.) are the most frequent
associates ; and Jeffrey pine (Pinus Jeffreyi Murr.) is a poor third.
A surprising fact observed at once on the upland west of the
mouth of Deadman Creek was the nearly exclusive presence of
large trees, juniper, fir, lodgepole and Jeffrey pine. Seedlings
and saplings were practically non-existent.

Several points deserve mention in connection with the reasons
why these junipers occupy such inhospitable sites and how they
manage to exist. (1) The junipers do not seem able to exist in
close association. From observation, however, it does not seem
likely that they crowd out other species. More probably the
latter cannot gain a foothold on certain locations. (2) The
extraordinary vitality of the junipers permits them to live on
what appears to be almost sterile granite or on coarse gravels
deficient in large quantities of mineral nutrients. (3) In addi-
tion, these junipers possess remarkable regenerative powers. An
injury must be serious indeed even to cripple a tree, much less
to destroy it. Injuries which are mortal to less hardy species,
such as pine and fir, or which shorten their lives to a marked

1937] GLOCK: WESTERN JUNIPER 23

degree, may only deform the juniper. New wood may overlap
a wound on the trunk so completely as to defy detection from the
exterior. Root buttresses and multiple trunks become so closely
compressed that the symmetry of the trunk is enhanced rather
than lessened. Stumps and cores showed bark and decayed
places where none would be expected from an exterior examina-
tion. Indeed, in some trees it was nearly impossible to secure
an uninjured core eleven inches long. (4) A tree remains alive
as long as a single root and branch function. It may be said
that the junipers die by inches. Many specimens were seen
which, save for one tuft of green, were gaunt gray skeletons.
So numerous are the partially dead branches that one is tempted
to say they are typical of all trees. The tops and sides of the
main branches are commonly devoid of bark, dead, and deeply
weathered. One-third or less of the girth of a branch, on the
under side, may support bark, beneath which lies active cambium.
One specimen showed only about 15 per cent of functioning
wood. As a consequence of growth restricted to the under side,
the branches are extremely eccentric, or hyponastic, and so much
so that the vertical diameter may be three to five times the hori-
zontal. (5) The ability to use soil moisture when it is available
is an important feature that will be taken up in the discussion on
the time of formation of the annual ring.

The western juniper merits distinction because of the size of
its trunk and because of the great age attained by its individuals.
Trees whose diameters range from four to eight feet are very
common. On the Kit Carson road several miles west of Kit Car-
son Pass, the Forestry Service has erected a sign reading ‘‘Cali-
fornia’s Largest Juniper” near a tree which is stated to be 31
feet 8 inches in circumference. However, since it is a triple-
trunked tree its age cannot be as great as this circumference
would indicate. Several large specimens were measured at a
height of five or six feet above ground on the upland west of the
mouth of Deadman Creek. One, whose trunk is a single shaft,
has a circumference of 27.5 feet. A second, triple-trunked
above, has a circumference of 31 feet.

As a species the western juniper ranks well above its com-
mon associates in the matter of longevity. Trees five and six
feet in diameter are not at all uncommon and those from which
core samples were taken gave evidence of being between 900
and 1000 years old. The longevity of the species certainly
equals that of the coast redwood and in a few instances rivals
that of the giant sequoia. Perhaps it is significant that the se-
_ quoia, the juniper, and the bald cypress of Oaxaca, Mexico, all
long-lived trees, belong to kindred families. These genera have
tremendous durability of their woods coupled with vigorous
tenacity of life and regenerative powers.

24 MADRONO [Vol. 4

The Bennett Juniper, a magnificent specimen, with a single
stately trunk approximately eighty feet high, is the giant of all
those so far observed by Mr. Clarence K. Bennett* of Hills-
borough, California, who has studied and hunted the juniper for
many years. It stands on the upland west of the mouth of Dead-
man Creek, a tributary of the Middle Fork of the Stanislaus
River. The dimensions of this large juniper are as follows:

Circumference:ateround | -ee 57 feet, 6.5 inches
Circumference six feet above ground .. 42 ” 9 >.
Greatest) diameter! ate ground ome: 21 ean
Average diameter five feet above ground 14 ” 2 a,

The diameter for working purposes may be taken as 13.5 to
14 feet.

It is highly appropriate that this monarch among junipers be
named “‘The Bennett Juniper,” not that Mr. Bennett was the first
to see it (hunters saw it previously), but because he was the first
man to take a lively sustained interest not only in the giant but
also in all junipers of its kind wherever he has been able to find
them in the high Sierra Nevada.

From the two or three stumps noted it was surmised that in
this region the species has yielded but little to the saw commer-
cially. Age, therefore, had to be determined wholly from cores
which were obtained from trees of different diameters. Usually
several cores, varying in length from six to fifteen inches, were
taken from each tree. In computing age, due allowance must be
made for increasing width of rings toward the center of the tree.
This increase, however, is not nearly so striking as in the pine.
All cases actually observed showed thinner rings at the center
than several inches outside the center and in no case was there a
uniform increase in width inward from the bark. Nevertheless,
ample allowance for the “‘age curve” was made in order to keep
estimates on the conservative side.

The age of the Bennett Juniper was computed in three ways,
in the first two of which synthetic trees were built up by the
substitution of progressively smaller trees for the inside of the
big tree. For the first trial, four trees from the same upland
which supports the big tree were united with five specimens from
the big tree itself. The four trees included four cores from an
8.5-foot tree, three cores from a 4.5-foot tree, one core from a
27-inch tree, and a section of a 6-inch tree. The rings in the
spaces between the cores and the next smaller tree were com-
puted by interpolation. In round numbers, the age of the Ben-
nett Juniper came out as 2900 years.

Entirely new material gave the basis for the second estimate.
Seven cores, the longest fifteen inches, were taken from the big

1 Clarence K. Bennett. The largest juniper? Sierra Club Bulletin 18:
115-116. 1933.

1937]

Puate VII.

GLOCK: WESTERN JUNIPER

Tue BENNETT JUNIPER (JUNIPERUS

Hook.).

OCCIDENTALIS

26 MADRONO [Vol. 4

juniper at about equal intervals around the trunk, save for the
northwest where there was some dead wood. These cores aver-
aged more than seven hundred rings to the first foot of radius.
Two trees from Charity Valley, one 8 feet and the other 5 feet in
diameter, were inserted for the interior of the large tree. The
first foot of the 8-foot tree had 476 rings and the first foot of the
5-foot tree had 510 rings. By computation and interpolation the
age of the Bennett Juniper came out to be 3250 years.

The third estimate of age was based on the ring counts in the
seven cores. Ring counts were plotted and the smooth curve
drawn through the plotted points was continued inward to the
center of the tree. The age so computed averaged 3000 years
or slightly more.

From the data at hand, a fair estimate, in round numbers, of
the age of the Bennett Juniper is 3000 years. There can be little
doubt that the tree is well over 2000 years old, since the outer
foot of the trunk averages seven hundred rings. The age curve
would have to be steeper than any observed in other trees in
order to hold the age below 2000 years. It may well be that the
age is distinctly more than 3000 years, but until the entire stump
is visible the exact age will not be known.

Two interesting problems that merit comment arose during
the study of the western juniper: (1) circuit uniformity; (2) time
of ring formation. |

Circuit Uniformity. Annual rings are said to possess circuit
uniformity if their relative thicknesses remain constant around
the entire circumference. A juniper 19 feet high and 6.5 inches
in diameter six feet above ground was felled on the upland west
of the mouth of Deadman Creek, about seventy-five yards from
the Bennett Juniper and up slope from it. None of the half
dozen sections taken at different heights shows a high degree of
circuit uniformity. In fact three or four rings show unusual
thickening at one place on the circumference and groups of sev-
eral other rings at other places. Few rings are uniformly thicker
or thinner than their adjacent neighbors. This being true, it is
to be expected that difficulty must attend the attempt to match
rings in different trees or in different radii of the same tree.
However, a few isolated cases of such matching, or cross-dating,
were detected at this elevation.

The lack of perfect uniformity in the western juniper resem-
bles somewhat that described by Shreve? for the Monterey pine
(Pinus radiata Don), save that in the case of this single juniper
vertical uniformity appeared to be better developed than circuit
uniformity. On the other hand, extensive and detailed studies
have shown remarkable uniformity in certain species, especially
the western yellow pine of the southern part of the Colorado

2 Forrest Shreve. The growth record in trees, Carnegie Inst. Wash. Pub.
No. 350, 91-116. 1924.

1937] GLOCK: WESTERN JUNIPER 27

Plateau. Is ring uniformity an environmental or a specific char-
acter? |

Time of Ring Formation. In the case of the 19-foot juniper,
on July 5, when snow had been off the ground less than two
weeks, the ring for 1936 was one-third to one-half the thickness
of the previous ring, or about equal in thickness to the average
ring. Tip growth already exceeded three inches. Cores taken
at the same time from junipers at the same locality and at Chip-
munk Flat showed rings for the current season of one-tenth to
more than one-half the average thickness of the last six rings.
A lodgepole pine had grown a ring more than three-fourths the
thickness of the average for the last six rings. Four cores from
Digger pines (Pinus Sabiniana Dougl.) which grew six miles
southwest of Sonora in the lower foothills gave evidence that
their seasonal growth was practically completed by July 6. In ©
at least two cases summer wood had been formed. Cores taken
from the Bennett Juniper on August 7 possessed what seemed to
be practically complete rings for 1936. The same was true for
the junipers from Charity Valley. A lodgepole pine from the
vicinity of the Bennett Juniper had its growth nearly completed
but neighboring Jeffrey pines were a trifle behind in their growth.
Lodgepole pines from Hope Valley had just begun the formation
of summer wood as had also western yellow pine from Walker
Canyon (6200 feet) near Sonora Pass Junction. .Two lodgepole
pines from Sonora Pass at 9650 feet elevation were beginning
the formation of summer wood. Judgment as to the complete-
ness of the annual ring was made not alone upon the amount of
wood in relation to previous rings but also upon the two follow-
ing criteria: diminution of cell size and deposition of dark ma-
terials on cell walls. In the Digger pines and one western yellow
pine from Walker Canyon dark color had appeared. Certainty
of the correct identification of cellular division for 1986 was
enhanced by cross-dating, that is, the matching of characteristic
rings from one tree to another.

In connection with the above observations three points have
considerable significance. (1) Growth must certainly begin in
the high-altitude trees here studied before the snow leaves the
ground. (2) The evidence indicates that a considerable part
(more than half in some cases) of the seasonal growth is com-
pleted within two weeks after the snow has melted. This applies
in particular to the locality of the Bennett Juniper on the west
end of East Flange Ridge. Where the water supply is unques-
tionably greater, as is the case for the trees at Chipmunk Flat,
there are indications that growth was less than at the previously
mentioned locality. Both Chipmunk Flat and East Flange Ridge
are so situated that they no doubt receive cold air drainage, but
the former much more so than the latter. (3) The time of maxi-
mum ring formation early in the season, before temperature has
reached a maximum and before water supply has diminished to a

28 MADRONO [Vol. 4

serious extent, has important climatic significance. If the obser-
vations so far made are valid, then the length of the so-called
growing season as based upon the interval during which tempera-
tures are supposed to be advantageous for growth has little influ-
ence on the thickness of the annual ring. Apparently the trees
must make their growth while water is available. If it is avail-
able all summer, growth continues for a longer time; but if water
is dependent upon winter precipitation and receives no replenish-
ment later, growth ceases sooner or later during the summer
according to the availability of the water. The Digger pine
from the foothills below Sonora was the most advanced in growth
of all trees from which core samples were taken on July 5 and 6.

Such evidence as has been obtained suggests that the time of
ring formation constitutes a problem which deserves study by
taking core samples periodically during an entire growing season
of trees from the lower forest border up to timber-line. The
presence in any particular area of trees which depend solely
upon winter precipitation has a climatic and ecologic significance
different from the presence of those which depend upon a sum-

mer rainy season.
Carnegie Institution of Washington,
Division of Plant Biology,
Tucson, Arizona,
September 4, 1936.

A NEW SPECIES OF CHAETOMORPHA FROM CHINA

NATHANIEL Lyon GARDNER

On a recent visit to the California Academy of Sciences in
Golden Gate Park, San Francisco, my attention was called by
Dr. F. M. McFarland, President of the Academy, to a small tor-
toise which had been recently asquired through the generosity
of Princess Olga Shahovski, who had brought it from China. Its
back was densely covered with “green hair,” so popularly desig-
nated. Such a symbiotic relationship, if such it may be called,
is of very rare occurrence in western America, if indeed it has
ever been observed and reported among our native species. How-
ever, it has been reported several times as occurring in the eastern
part of the United States, and apparently is of common occur-
rence in parts of China and Japan. Wang (5) states that the
“green haired tortoise” has been a well known animal in the
provinces of Changshu, Kiangsu, and in the northern part of Yu-
shan. In China the “green hair’ grows on relatively small spe-
cies of tortoise. These of course are not of the basking kind,
and on account of the popular interest which they create in
domestic aquaria they are commercialized to considerable extent.

The specimen in the public aquarium at the California Acad-
emy rarely fails to excite the interest and curiosity of visitors,
often calling forth strange expressions indicative of total igno-

1937] GARDNER: CHAETOMORPHA 29

rance as to the real nature of the organisms, the visitors never
having previously seen nor read of similar or like phenomena.
It thus seems desirable to publish a few brief statements con-
cerning them. The so-called “green hair’ on the tortoise is of
course not a part of the animal but is a distinct organism belong-
ing to the plant kingdom, and to a group known as Chloro-
phyceae, or green algae. The determination of the species, how-
ever, is not in this case such a simple matter as it would seem at
a glance.

A brief search through the literature revealed the fact that
several papers have been written concerning the occurrence of
species of green algae growing on the backs of the tortoise.

Collins (1) reports Chaetomorpha chelonum growing on the
back of Chrysemys marginata and Aromochelys odorata found in
Michigan. Evermann and Clark (2) later reported the same
species growing on Chrysemys marginata in Indiana, and still later
Tiffany (4) found it growing on Chrysemys marginata belli in
Iowa. Hoffmann and Tilden (3) described a species growing
on Chelydra serpentina found in Minnesota, which they considered
related to Collins’ Chaetomorpha chelonum but sufficiently different
to warrant the establishment of a new genus, Basicladia. Various
other authors in this country and in Europe have incidentally
mentioned, in connection with their study of the tortoise, the
occurrence of green algae growing on the backs of these animals
but without any attempt to name the species to which the algae
belong. Yendo (6) found plants in Japan growing on Clemmia
japonica which he described as a new variety of Chaetomorpha
chelonum Collins. The most recent paper with which the writer
is familiar, dealing with the subject, is by Wang (5), who seems
to have made the most exhaustive study of species growing in
China. He states that he has examined four hundred and seventy-
five specimens of the tortoise Geoclemys reevesii. He reports two
species in this paper, viz., Basicladia crassa Hoffmann and Tilden,
and Cladophora glomerata var. nana Wang.

The green alga on the “green haired tortoise” with which this
paper is especially concerned is growing on a small tortoise,
Ocadia sinensis. Only one living specimen is available for this
study and this has been in captivity at the Academy some four
or five months. Apparently the reproductive stage of the alga
has passed. The presence of many empty segments, each with a
specialized pore for the escape of reproductive cells, gives ample
evidence of the fact that the plants are normal, but there seem
to be no more fertile segments, hence I am unable to report on
the character of the reproductive cells. Strange to note, not one
of the authors dealing with these plants has described the repro-
ductive cells, not even Mr. Wang, who examined the large num-
ber of specimens mentioned above. Knowledge of the character
of these cells is of very great importance in connection with this
species of plant, since it has morphological characters linking it

30 MADRONO [Vol. 4

closely with at least four different genera, viz., Chaetomorpha
Kuetz., Rhizoclonium Kuetz., Basicladia Hoffmann and Tilden, and
Cladophora Kuetz. The principal distinguishing characters of
these genera may be noted as follows:

CHAETOMORPHA

Fronds rigid, septate coenocytes, unbranched, always attached
in the juvenile stage by more or less branched, nonseptate rhizoids
from the basal segment and whose walls become thickened and
whose contents disappear in age; asexual reproduction by bicili-
ated zoospores.

RHIZOCLONIUM

Fronds septate coenocytes, usually narrower and more flaccid,
not attached, unbranched or producing few to many short, irregu-
lar, septate or nonseptate, rhizoidal branches from any of the seg-
ments; asexual reproduction absent.

BASICLADIA

Fronds consisting of numerous, multicellular, erect, somewhat
rigid, sparingly branched, more or less cylindrical filaments aris-
ing from creeping rhizome-like filaments which are fastened to
the substratum by hold-fasts having free or coalesced branches;

. reproduction by zoospores (?). (An extract from the origi-
nal description of the genus.)

CLADOPHORA

Fronds septate coenocytes, rigid to flaccid, always moderately
to profusely branched, always attached in the juvenile stage by
the discoidal basal segment, later often producing more or less
abundant septate, rhizoidal filaments from the lower parts;
asexual reproduction by four-ciliated zoospores.

The following is a diagnosis of the species under considera-
tion:

Chaetomorpha sinensis sp. nov.’ Erect fronds producing
tangled, more or less rope-like masses, very sparsely branched,
the branches being either similar to the main axes or rhizoidal in
character, attached by relatively extensive, prostrate, irregular,

Chaetomorpha sinensis sp. nov.

1 Frondibus erectis sparsissime ramosis structuram compositam funibus
irretis aliquantum similis producentibus; ramis axis principalibus similibus vel
rhizoideis, filamentis amplis, septatis, prostratis irregularibus plus minusve
ramosis adjunctis; filamentis substrato superficiebus nec hapteris praecipuis
nec ramis rhizoideis adhaerentibus; ramis filamenta secundaria, 8-12 cm. alta,
60-95 wu diametro, filamentis primariis similibus aliquando emittentibus; fila-
mentorum erectorum segementis maximam partem etiam in generatione cylin-
driciis, sed interdum leviter doliiformibus longitudine maxime variabilibus
basim 2 mm. superne diametro 3-4-plo longioribus; filamentorum parietibus
comparate tenuis et tenacibus, hyalinis demum plus minusve lamellatis; chro-
matophoris densis; pyrenoidibus parvis, numerosis; cellularis generationis
foraminibus caminis parvis similibus in regione equatoriale segmentorum
fertilium discedentibus; cellularum generationis natura ignota.

1937] GARDNER: CHAETOMORPHA 31

2

Pruate VIII. CuarromorrHa SINENSIS GARDNER. Filaments X100. 7. Frag-
ment of a creeping filament, with one small juvenile erect filament, to the left;
one segment of a normal erect filament showing laminated cell wall, in the center;
and one dead erect filament with three segments, to the right. 2. Single seg-
ment of a medium sized filament showing numerous pyrenoids embedded in
normal chromatophores. 3. Two wider, shorter segments. 4 Fragment of a
normal filament showing empty segments with the chimney-like pores through
which the reproductive cells have escaped.

°

32 MADRONO [Vol. 4

more or less branched, septate filaments adhering to the sub-
stratum by their surface and not by specialized hapteres or rhi-
zoidal branches, and at times giving rise to secondary filaments
similar to the primary ones, 8-12 cm. high, 60—95 y diam.; seg-
ments of erect filaments mostly cylindrical, even in reproduction,
but occasionally slightly dolioform, very variable in length, up to
2mm. at the base to 3—4 times the diameter in the upper parts;
walls relatively thin and tough, hyaline, more or less lamellate in
age; chromatophore dense; pyrenoids small and numerous; re-
productive cells escaping through specialized chimney-like open-
ings formed in the equatorial region of the fertile segments; the
nature of these reproductive cells not determined.

Growing principally on the back of a fresh-water tortoise,
Ocadia sinensis, brought from Kiangsu Province, China. Type,
Herbarium of the University of California no. 543979.

By comparison of the above diagnosis with the principal dis-
tinguishing characters of the four genera mentioned above, it
may be seen that it has overlapping morphological characters.
It is relatively harsh and rigid, like Chaetomorpha; an occasional
true branch on the erect filaments links it with Cladophora; the
more abundant rhizoidal filaments from the erect fronds are
homologous with the same in Rhizoclonium and similar to those
found at times in Cladophora. The extensive prostrate, attaching
filaments are similar to those upon which, partially, the genus
Basicladia was established. They differ, however, from these in
the method of attachment as stated in the diagnosis of that genus.
It is therefore essential to know the character of the reproductive
cells before determining positively to which genus it should be
associated. Are the reproductive cells gametes or zoospores, and
if the latter, are they biciliated or quadriciliated? For the pres-
ent I am placing it as a new and eccentric species of the genus
Chaetomorpha, with a more extensive attaching system than the
general run of species, and with an extremely occasional true
branch on a very few plants.

University of California,
Berkeley, June 12, 1936.

LirerATURE CITED

1. Corrins, F. S. Some new green algae. Rhodora 9: 197-202. 1907.

2. EvermMann, B. W., and Crarx, H. W. The turtles and batrachians of the
Lake Maxinkuckee region. Proc. Ind. Acad. Sci. 1916. 472-518. 1917.

3. HorrMann, W. E., and Tinpen, J. E. Basicladia: a new genus of Clado-
phoraceae. Bot. Gaz. 89: 374-384. 1930.

4. Tirrany, L. H. The filamentous algae of northwestern Iowa, with special
reference to the Oedogoniaceae. Trans. Amer. Mic. Soc. 45: 69-132.
1926. :

5. Wane, C. C. Algae growing on the pond tortoise. Contrib. Biol. Lab. Sci.
Soc. China 10: 4-9. 1935.

6. YrenNpbo, K. Nov. Alg. Japon. Decas 1-111. Bot. Mag. Tokyo 34: 1-12.
1920.

1937] GANDER: FLORA OF SAN DIEGO COUNTY 33

NOTES ON THE FLORA OF SAN DIEGO COUNTY,
CALIFORNIA

Frank F. GAnper

Extensions of Range

The extensions of known ranges of the species listed below
seem worthy of record. Except where otherwise designated, col-
lections were made by the writer. Numbers in parentheses refer
to specimens in the herbarium of the San Diego Natural History
Museum.

SmILAcINA sTELLATA (L.) Desf. Collected in a low, wet place
near Cuyamaca Lake at an elevation of about 4600 feet, in April,
1934, by Miss Eleanor C. Layman (722) ; not previously reported
from this county.

CHoRIZANTHE OrcutTiana Parry. Heretofore known only from
the type locality on Point Loma, this species was found on
Kearny Mesa north of San Diego, about nine miles from Point
Loma, March 18, 1935 (10604). The somewhat similar C. poly-
gonoides Torr. and Gray was found in the same vicinity.

SAXIFRAGA CALIFORNICA Greene. This species was collected in
San Diego by Daniel Cleveland in April, 1874 (8384), and in
March, 1875 (8882). In recent years it has been collected in
San Diego and at the Bear Valley School near Escondido. Previ-
ously it was not recorded south of the Santa Ana Mountains of
Orange and Riverside counties.

MENTZELIA MICRANTHA (Hook. and Arn.) Torr. and Gray.
This blazing star was found on Otay Mountain, just north of the
Mexican boundary, June 26, 1935 (11696).

JUSSIAEA CALIFoRNICA (Wats.) Jepson. San Luis Rey River,
near San Luis Rey Mission, July 10, 1985 (12057).

MyrioPHYLLUM EXALBESCENS Fernald. Collected in Lake Mur-
ray, July 27, 1985 (12083); probably introduced. The plants
were in full flower.

VENEGASIA CARPESIOIDES DC. Although reported from San
Diego County, there are apparently no definite records of locality
for this species. It was collected in Moosa Canyon, north of
Escondido, March 18, 1935 (10571), and in near-by Cole Canyon
on September 18, 1935 (12150).

Records of Alien Plants

Most of the alien species listed below have not previously
been reported from San Diego County, and one (Echium) is here
reported from California for the first time.

THLASPI ARVENSE L. Collected in Balboa Park, San Diego, by
Miss Fidella G. Woodcock; date not given (2009).

Coronopus pipymus (L.) Smith. A number of specimens were

collected on a parking lot in Balboa Park, San Diego, March 21,
1936 (14122).

34 MADRONO [Vol. 4

Resepa ALBA L. Several plants were found growing along the
bank of the San Luis Rey River across from Pala on April 25,
1935 (11302). |

Gaura stnuatTa Nutt. Three stations for this species are rep-
resented by specimens in our herbarium, but it has also been
observed at several additional localities. It was first reported in
July, 1980, when W. V. Shear collected specimens at Carlsbad
(8355). It was found growing along the highway two miles north
of Lake Hodges on March 6, 1934 (3354), and was thoroughly
established in a field at Santa Ysabel on July 10, 1935 (11940).

Ecuium PLANTAGINEUM L. Not previously reported in Cali-
fornia, this species was found growing abundantly in a meadow
and around a spring by the roadside near DeLuz, May 1, 1935
(11821).

Lycium Hauimirotium Mill. Collected by Charles F. Harbison
in the bed of the Tijuana River about two miles from its mouth,

August 19, 1934.
Natural History Museum, Balboa Park,
San Diego, California,
January 30, 1936.

NEW RECORDS OF VASCULAR PLANTS IN
WASHINGTON

GrorGE NEVILLE JONES

Even in a region which has been as well botanized as Wash-
ington, there are many species of vascular plants whose known
occurrence rests upon a single collection or a more or less definite
statement. The following notes record eleven species of vascular
plants not hitherto ascribed to this state. These include six recent
immigrants and five indigenous species. The records are based
on specimens in the Herbarium of the University of Washington.

ANEMONE Lupoviciana Nutt. Gen. Am. Pl. 2: 20. 1818. This
species is abundant on the prairies and plains east of the Rocky
Mountains. Although recorded by Rydberg (8, p. 288) as occur-
ring in Washington, until very recently there have been no speci-
mens in local herbaria to substantiate the record. This spring,
however, a number of plants of this anemone were sent to the
University of Washington to be identified. The collection data
are as follows:

Chelan County: foothills near Wenatchee, May 15, 19386,
Doris Mullen.

Crematis Viratsa L. Sp. Pl. 544. 1753. During the last
thirty years or so this European species of Clematis has become
well established in various localities in western Washington. It
is quite common at Seattle and Tacoma. It is sometimes mis-
taken for the indigenous C. ligusticifolia Nutt. of the Upper So-

1937] JONES: WASHINGTON PLANT RECORDS 35

noran zone east of the Cascade Mountains, from which it differs
in its perfect flowers, more densely pubescent sepals, and less-
notched leaflets. The following Washington specimens are at
hand:

San Juan County: Roche Harbor, July 11, 1904, A. S. Pope;
King County: Seattle, Jones 8567; Fall City, Jones 9426; Pierce
County: Tacoma, Jones, December 28, 1927.

PoLANIsIA TRACHYSPERMA Torr. and Gray, Fl. N. Am. 1: 669.
1840. This species was noted (as P. graveolens) in the report of
the Wilkes Expedition (5, p, 235) as occurring along the Walla
Walla River in what is now the state of Washington. Piper dis-
missed it from the “Flora of Washington” (2, p. 307) with the
remark that ‘There are no herbarium specimens of this plant
from Washington to justify its inclusion in the flora.” The
record of the following collection will serve to establish the
actual occurrence of the plant in Washington at the present time:

Walla Walla County: gravelly bank of the Columbia River
near the mouth of the Walla Walla River, June 22, 1934, Jones
5085. The specimens show both flowers and fruit. The general
range of this species is from British Columbia to Texas and as
far east as Iowa and Saskatchewan.

Rosa rucosa Thunb. Fl. Jap. 218. 1784. At the time of the
publication of my recent paper on the species and varieties of
Rosa (1), R. rugosa was known to me from only one locality in
Washington. I concluded, therefore, that this species was only
an incidental “garden escape” and not of sufficient importance to
be included in the rose flora of the state. Since that time, how-
ever, several additional reports of this rose have come to my
attention. Apparently the species is becoming well established
in various localities in western Washington, especially near the
seashore. The following collections are at hand:

Island County: Useless Bay, Whidby Island, June 2, 1934,
Jones 6133. Kitsap County: Restoration Point, Bainbridge
Island, April 19, 1936, Jones 8734.

VioLa LANcEoLATA L. Sp. Pl. 9384. 17538. The recent dis-
covery of this violet in western Washington extends the known
range of the species a thousand miles to the westward. It was
previously known to range from Nova Scotia to Florida and
westward to Minnesota, Nebraska, and Mississippi (4, p. 554).
A report of such a remarkable extension of range is, I am aware,
not above suspicion; but there is not the slightest doubt as to
correctness of the identification of the plants from western Wash-
ington. That they could be adventive here is possible, though
from the ecological evidence, highly improbable. They occur in
abundance in several rather widely separated localities, in habi-
tats apparently never disturbed by human activity. That they
have not been previously discovered here is probably due to the
fact that they are inconspicuous, except when in flower, and

36 MADRONO [Vol. 4

then most likely have been mistaken for V. pallens (Banks)
Brainerd, or even for V. palustris L., both common species in the
vicinity. In western Washington, Viola lanceolata may be a re-
lictual species; at least its habitat is on the outwash plains near
the southern limit of the Pleistocene glaciation.

Pierce County: in marshy ground between Tacoma and Roy,
April 27, 1936, Jones 8774.

Lepum GLanpu.tosum Nutt. Trans. Am. Phil. Soc. 8: 270. 1843.
This species ranges from British Columbia to California and east-
ward to Wyoming. It is known from several localities in Okano-
gan, Chelan, and Kittitas counties in eastern Washington, but has
been unknown, until very recently, from the western side of the
Cascade Mountains. This Labrador tea may be distinguished
from the common L. groenlandicum Oeder by its oblong or oval
plane-margined leaves which are green and glabrous on both
sides. It has been collected at the following locality in western
Washington:

King County: Delta Lake, J. M. Broadbent, August 5, 1935.
Mr. Broadbent, a student of botany at the University of Wash-
ington, reports the shrub to be fairly abundant on moist ledges
on the mountainside near the outlet of the lake. Prominent
among associated species were Rhododendron albiflorum, Saliz sp.,
Vaccinium macrophyllum.

SWERTIA PERENNIS L. Sp. Pl. 226. 1753. Not hitherto col-
lected in Washington, this gentianaceous perennial is known to
occur in British Columbia and Alaska, and in the Wallowa Moun-
tains in northeastern Oregon (August 25, 1898, Cusick 2100).
The first record of the collection of this plant in Washington is
as follows:

Snohomish County: on talus, Twin Lakes, altitude 4000 feet,
(in full bloom) September 2, 1935, J. M. Broadbent.

SOLANUM RostRATUM Dunal, Hist. Solan. 234. 18138. Adven-
tive in Benton County: Benton City, September 16, 1936, Harold
Stringer.

Gaxium verum L. Sp. Pl. 107. 1753. Long since naturalized
in the eastern half of North America, this Eurasian species can
now be listed as adventive in western Washington on the basis of
the following collections:

King County: weed in lawn, Seattle, October 20, 1932, Jones
4814; October 10, 1934, no. 6117. This is the only uncultivated
species of Galium in Washington with yellow flowers. The plants
are perennial, with linear, deflexed leaves in whorls of six or
eight.

Veronica CHamarprys L. Sp. Pl. 138. 1753. This is a per-
ennial with the stems pubescent in two lines, the leaves subses-
sile, cordate, incisely crenate, and the flowers blue, 4-6 mm.
broad, appearing in May. The plant has been noted in several

1937] REVIEWS 37

places in Seattle, where it occurs chiefly as a weed in lawns, in-
troduced, probably, with grass seed. It is represented by the
following collection:

King County: Seattle, weed in lawn on the campus of the
University of Washington, May 2, 1933, Jones 4374.

Taraxacum LaEviecatum (Willd.) DC. Cat. Hort. Monsp. 149.
1813. Readily distinguishable from the more common T’. offici-
nale Weber by its bright reddish achenes and the leaves dissected
almost to the midvein, this species is now established in several
places in Washington. Probably it is frequently mistaken for T.
officinale and for that reason is rarely collected.

King County: Seattle, March 17, 1934, Jones 8721.

University of Washington Herbarium,

Seattle, Washington,
October 20, 1936.

LiIvERATURE CITED

1. Jones, G. N. The Washington species and varieties of Rosa. Madrojfio 3:
120-135. 1935.

Piper, C. V. Flora of the state of Washington. Contr. U. S. Nat. Herb.
11: 1-637. 1906.

Rypserc, P. A. Flora of the Rocky Mountains. 1917.

—————._ Flora of the prairies and plains. 1932.

Torrey, J. U.S. Exploring Expedition 17 (Wilkes Expedition) : 205-514.
1874,

REVIEWS

The Genus Arabis L. in the Pacific Northwest. By Rerep C.
Roxtuns. Research Studies of the State College of Washington,
Volume IV, Number 1. Pp. 52, with 15 figures. Pullman, Wash-
ington, 1986. $.50.

Perhaps no western group of flowering plants has been in a
more confused state than has the genus Arabis, and all system-
atists will be glad to know that an attempt has been, and is being,
made to set this wing of the cruciferous house in order. More
than sixty species have been described from, or attributed to,
the states of Washington, Oregon and Idaho—Greene, Howell
and Piper each having added his quota. Mr. Rollins has examined
these critically, submitting them to the important test of geo-
graphical significance. After four species have been excluded,
only twenty-one species and eleven varieties survive his scrutiny.
The classical criterion of the uniseriate versus the biseriate con-
dition of the seeds is examined and explained by observation of
ontogenetic development. The kind of pubescence, when pres-
ent, is found to have considerable diagnostic value, whereas
degree of pubescence is significant only within broad limits.

Fifteen nearly full page original line drawings by Mr. Rollins
depict representative and usually confused species, and two maps
graphically show the geographical basis of this study. A num-
ber of subspecific populations, which have previously posed as

38 MADRONO | [Vol. 4

species, have been reduced to varietal rank, making a total of
seven new combinations.

It is gratifying to know that through an assistantship at Gray
Herbarium Mr. Rollins will be enabled to expand his study to
include Arabis of western North America.—Lincotn ConsTANce.

Handbook of Northwest Flowering Plants. By HELen M. GILKeEy.
Pp. 407. Metropolitan Press, Portland, Oregon, 19386. $2.25.

The author describes her book as “‘an illustrated hand-book
of the more conspicuous plants of the Northwest,” designed espe-
cially for beginning students and interested laymen. The area
covered is roughly that which Piper termed “the Vancouver
Strip,” that is, the region from the summit of the Cascades to the
Pacific, and from the northern boundary of the California flora
to northern Washington. The author considers technical ques-
tions of nomenclature and specific concept to be beyond the
scope, and aside from the main purpose, of this volume. Insofar
as possible, Dr. Gilkey has avoided technical terminology in keys
and descriptions, and has omitted many of the groups of especial
difficulty or of less interest to amateur flower lovers.

The illustrations constitute a most distinctive feature of the
flora, text figures of more than three hundred species making the
book invaluable for students of limited experience. The family
and generic lines are closely in accordance with Jepson’s ““Manual
of the Flowering Plants of California”; the nomenclature is con-
servative; rare and doubtful species are, for the most part,
excluded or mentioned only briefly. There are no new names
published in this work.

Despite its modest pretensions, the new handbook is a worth-
while effort to fill, in part, the void left by the exhausting of avail-
able copies of Piper and Beattie, “Flora of the Northwest Coast.”’
—Lincoin Constance.

The Early Flowering of Plants in Lane County, Oregon, in 1934.
By Lovis F. Henperson. University of Oregon Monographs,
Studies in Botany, Number 1. Pp. 16. Eugene, Oregon, June,
1986. $.25.

The precocious season of 1934 caused much speculation as to
just how premature that spring and summer actually were. It
has remained for Professor Henderson, speaking from fifty years’
experience with the flora of the Northwest, to subject these con-
jectures to a scientific analysis. For each of 165 species the
earliest date of blooming in 1934 was compared with the average
date of blooming compiled from collections made during the last
four or five decades. His general conclusions are that most early
spring plants bloomed two months in advance of their usual time;
trees, 1 month and 7 days; water plants, 1 month; Hudsonian-
Alpine zone plants, 1 month and 2 days; bog plants, 1 month, 7
days; early spring herbs, 2 to 33 months; late summer flowers,

1937] REVIEWS 39

1 month to 1 day earlier, or even later than the average.—Lin-
COLN CONSTANCE.

A Botanical Survey of the Olympic Peninsula, Washington.
By Grorce Nevitte Jones. University of Washington Publica-
tions in Biology, Volume V. Pp. 288, with 9 plates. Seattle,
June 25, 1986. $2.00.

In this excellent study, Mr. Jones has produced the most
thorough and successful treatment which any portion of the
Pacific Northwest has received in the last thirty years. The
Olympic Peninsula comprises some four thousand square miles,
and contains the Washington expression of the Coast Ranges, the
Olympic Mountains, which rise from sea-level to a height of
nearly eight thousand feet and are completely isolated, by water
or lowlands, from the neighboring Cascades and Oregon Coast
Ranges.

Ever since Menzies collected briefly along this coast in 1792,
the Olympics have been a prized ground for many collectors, in-
cluding Henderson, Piper, Flett, Heller, Elmer, Thompson and
many others. Several lists of species have appeared from time
to time, beginning with Henderson’s’, and all the known species
were included by Piper in his floras, but no prior attempt has
been made to consider the area in question as a distinct vegeta-
tional and floristic unit.

Not the least satisfactory portion of the present paper is the
introductory synecological and distributional discussion, modeled
after that of Piper’s Flora of Washington, and comprising nearly
one-fourth of the volume. Jones has made an original contribu-
tion in attempting a correlation of the “‘life-zone concept” of
Merriam with Raunkiaer’s “biological spectrum.” It will be re-
called that Raunkiaer grouped plants into “life-forms,” depend-
ing upon the mode of resistance of species to the critical period
of the life cycle, and gauged by the position of the perennating
organs. He defined the following groups: phanerophytes,
chamaephytes, hemicryptophytes, cryptophytes and therophytes.
Jones finds that the temperature and moisture conditions charac-
terizing a given biotic belt are clearly indicated, in the main, by
the definite proportional representation of these five different
life-forms. The Arctic-Alpine zone, for example, lacks phanero-
phytes and therophytes, and has 21 per cent chamaephytes, 9 per
cent cryptophytes and 69 per cent hemicryptophytes, whereas
the neighboring Hudsonian zone exhibits the following propor-
tions: 9 per cent phanerophytes, 10 per cent chamaephytes, 67
per cent hemicryptophytes, 13 per cent cryptophytes and 1 per
cent therophytes.

According to the author: “The significance of the Raunkiaer
system of life-forms as applied to the flora of the Olympic Penin-

1 Henderson, L. F. Flora of the Olympics. Zoe 2: 253-295. 1891.

40 MADRONO [Vol. 4

sula is threefold. By means of this system a statistical analysis
can be made of the flora of the whole region, or of the flora of
each of the four life-zones taken separately. By whichever
method used the results are comparable with the flora of other
regions, and a simple but biologically sound summary of the
phytoclimate thus can be obtained. As applied to Merriam’s
life-zones, the Raunkiaer system yields corroborative data. Mer-
riam was concerned chiefly with the factors of the climate which
are effective during the season of growth and reproduction,
whereas the Raunkiaer system is based on the adjustment of
plants to the unfavorable, which is usually the dormant, season.
By the application of both of these systems, a much clearer char-
acterization of the life-zones or climatic formations may be
obtained.”

The outstanding characteristics of the region are its isolation
and the wealth of climatic and altitudinal diversity which it mani-
fests. The peaks of the range afford a serious obstacle to water-
laden clouds, so that the 150-inch rainfall on the coast dwindles
to fifteen inches at irrigated Sequim, on the northeast. It is not
surprising, then, that the area presents a peculiar and interesting
flora. Although 75 per cent of the flora is that of the adjacent
Cascades, many Cascadian plants have not yet been found here,
and many species extend their ranges northward or southward to
the Olympic Peninsula, without occurring west of Hood Canal.
Of especial interest is the occurrence of a small group of en-
demics, probably relicts, which are almost exclusively confined
to unglaciated areas of high altitude, and which make up about
2 per cent of the total flora.

Over one thousand species and varieties are recognized for
the Olympic Peninsula, which is over one-third of the known
flora of Washington. The author exhibits laudable restraint in
proposing only four new species, three new varieties, one new
form and twenty-one new combinations, many of the latter being
worthwhile reductions.

By his comprehensive study of the vegetation and the distri-
bution of its species, with a consideration of the factors involved,
in addition to the annotated catalogue of vascular plants (pro-
vided with keys and citations of specimens), Jones has produced
a highly important piece of work. The prevalent attitude of the
past has been to consider the whole region only as a promising
field for the discovery of novelties. Similar attacks upon other
mountainous areas in the Northwest will afford opportunity for
significant comparisons with the flora of the Olympics. Now that
Jones’ study has revealed the necessity for more detailed investi-
gations of the Cascades before any very satisfactory compari-
sons can be made between the two ranges, it is to be hoped that
he will find time to give us a comparable study of Mount Rainier
—a project which he has long had in mind.—Lincoin Constance.

Complete Your Files!

MADRONO

A West American Journal of Botany

“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges.

Volume I, 1916-1929. . . $5.00
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ia

BOTANY

we

Contents

Elzada

»
Sy

(FORNIA Prace Names Usep spy DANIEL CLEVELAND ...

at Lime and Green Streets, Lancaster,
Pennsylvania

‘April, 1937

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asrams, Stanford University, California.

Dr. Lincotn Constance, State College of Washington, Pullman, Washington.
Dr. H. F. Coretanp, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Dr. P. A. Munz, Pomono College, Claremont, California.

Business Manager—Dr. Davin D. Kreck
Lime and Green Sts., Lancaster, Pennsylvania
or
Carnegie Institution of Washington
Stanford University, California

Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

Established 1916. Published quarterly. Subscription Price $2.50 per year.
Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17,
complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num-
bers $0.75.

Papers up to 15 or 20 pages are acceptable. Longer contributions may
be accepted if the excess costs of printing and illustration are borne by the
contributor. Articles may be submitted to any member of the editorial board
and if accepted, are published in the order of receipt. However, accepted
manuscripts may be included in the forthcoming issue provided that the
contributor pay the cost of the pages added to the issue to accommodate his
article. Reprints of any article are furnished at a cost of 4 pages, 50 copies
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at $1.50 per hundred. Reprints should be ordered when page proofs are
returned.

Published at Lime and Green Streets, Lancaster,
Pennsylvania, for the

CALIFORNIA BOTANICAL SOCIETY

President: Dr. F. W. Foxworthy, Berkeley, California. First Vice-Presi-
dent: Dr. Ira L. Wiggins, Department of Botany, Stanford University. Sec-
ond Vice-President: Dr. Philip A. Munz, Department of Botany, Pomona
College, Claremont, California. Treasurer: Dr. David D. Keck, Carnegie In-
stitution of Washington, Stanford University. Secretary: Miss Ethel Crum,
4004 Life Sciences Building, University of California, Berkeley.

Annual membership dues of the California Botanical Society are $2.00,
$1.50 of which is for a year’s subscription to Madrofio. Dues should be
remitted to the Treasurer. General correspondence and applications for
membership should be addressed to the Secretary.

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 4

VEGETATIONAL SURVEY OF THE LOWER
RIO GRANDE VALLEY, TEXAS?*

Evzapa U. CLover
BoTANICAL EXPLORATION

- Southern Texas has been known to botanists since 1826. The
earliest botanist was the Swiss, Dr. Luis Berlandier (4). He was
sent as naturalist of the Mier Terdn Expedition or Comision de
Limites by the Mexican Government to determine the character
of the country along the proposed United States and Mexican
boundary in 1828. Thomas Drummond, upon seeing a set of
Berlandier’s collections, was impressed by the vegetation of the
region and while on a collecting tour to America went to Texas
in 1838-1834 and spent some time collecting in the vicinity of
Galveston Island (38). W. J. Hooker (26) published notes on
this expedition. ,

Ferdinand Lindheimer began collecting in Texas in 1836 but

because of conditions in the early days of the Republic did no
extensive work until 1842. Dr. George Engelmann, a friend and
German schoolmate of Lindheimer, suggested to Asa Gray that
they take the burden of classification and distribution off his
hands, permitting him to devote his entire time to field work.
Lindheimer contributed much to the botanical knowledge of the
state and well deserves the title “Father of Texas Botany.” His
plants were described by Gray and Engelmann (18, 22).
In 1854 the work of making a United States-Mexican Bound-
ary Survey was begun. Major William H. Emory (17) was
placed in charge of this project. Vegetational and geological
reports of this survey by Dr. C. C. Parry and Assistant Arthur
Schott give interesting information on the Lower Rio Grande
region. Charles Wright, who, in connection with the movement
of troops to western forts, had made his first collections of plants
in Texas in 1847 to 1848, was in 1851 sent as botanist and sur-
veyor on the United States-Mexican Boundary Survey Commis-
sion from the Rio Grande to the Pacific, under Emory. Collec-
tions of plants (exclusive of the Cactaceae) made during this
survey were determined and distributed by John Torrey (34).
George Engelmann (19) determined the Cactaceae.

John M. Coulter (13) published on the plants of an expedi-
tion to the region of the Rio Grande made in 1887: “Mr. G. C.
Nealley was engaged by the Division of Botany (U. S. Depart-
ment of Agriculture) to make collections of plants during 1887,
1888, and 1889 in the more unexplored parts of Texas, chiefly in

1 Papers from the Department of Botany and the Botanical Gardens of
the University of Michigan, No. 625.

ManroNo, Vol. 4, pp. 41-72, April 7, 1937.

APR 1 6 1937

42 MADRONO [Vol. 4

the counties bordering the Rio Grande. It was hoped that many
of the rarer plants of the Mexican Boundary Survey and the
early collections would be rediscovered, that additional Mexican
types would be found to be members of our flora, and that species
new to science would be brought to light.” A large and valuable
collection was made on this expedition.

Valéry Havard (25) made extensive collections along the
valley of the Rio Grande and in the adjacent territory. His re-
port on this work really marks an important step in Texas botany
from pure description to a study of plants in relation to their
environment, ecology, and pathology. An account of Havard’s
work is given by Winkler (38).

C. Hart Merriam (28) in his much discussed “Life Zones and
Crop Zones of the United States,’ by mapping the Brownsville
region as “Tropical” called attention of botanists to the special
interest that the region might hold for them. Cultivated as well
as wild plants of the Rio Grande Valley are listed in the paper.

In an account of changes of vegetation in the south Texas
prairies, O. F. Cook (11) concluded that elimination of prairie
fires as a result of intensive grazing is the principal cause of the
spread of chaparral vegetation over much of the coastal prairie.

Dr. J. N. Rose made investigations in the Lower Rio Grande
Valley in connection with his studies in the family Cactaceae.
The great monograph of this family by Britton and Rose (7)
incorporates many notes and photographs by Robert Runyon of
Brownsville, an amateur botanist who has contributed valuable
notes on distribution as well as on new species. He was co-
author of “Texas Cacti’? (80) and published also an article in
Desert, May, 1936, on “‘Cacti of the Lower Rio Grande Valley.”
Another student of the flora of this region is Father Chateau of
Mission, Texas. ,

In this brief summary many collectors and authors who have
contributed to the knowledge of the botany of the Lower Rio
Grande have had to be neglected. However, the reader who has
a particular interest in the region may turn to Charles H. Wink-
ler’s account (88) of botanical investigations in Texas. He
includes an annotated list of 121 publications.

CLIMATE

The temperature variation in this region is not great. At Fort
Ringgold it averages by months from 57.7° F. in January to
88.7° F. in July; and at Brownsville, from 59.1° F. in January to
83.7° F. in August. These figures were compiled from statistics
of the U. S. Weather Bureau over a period of about forty-seven
years (85). Killing frosts are rare, but frequent enough to
make the commercial growing of bananas and other tropical
fruits impossible.

43

VEGETATION OF RIO GRANDE VALLEY

CLOVER

1937]

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Average monthly rainfall at Fort Ringgold varies from 0.29
inches in February to 3.13 inches in September, with a total of
17.46 inches annually; at Brownsville, from 1.27 inches in March
to 5.62 inches in September, with a total of 26.89 inches annually.
This record was made over a period of thirty-nine years at
Brownsville and thirty-seven years at Fort Ringgold (35).

The weather is usually clear with a brilliant sun; but in June
and September there are such violent thunder storms that the
usually dry, deep arroyos in Starr and Zapata counties become
impassable torrents. The prevailing wind in the Lower Rio
Grande Valley is from the southeast. It blows almost steadily in
the summer but decreases during the winter. It is of the mon-
soon type. ‘“‘Northers” occur with varying frequency from late
October until March, causing a very sharp drop in temperature
within a short space of time. They are called “dry” or “wet”
northers, depending upon whether or not the wind is accompanied
by precipitation. Since rainfall in this region is so irregular,
farming as a rule is very uncertain in unirrigated sections. How-
ever, cotton raised by dry farming methods seems to be quite suc-
cessful because the plants are less injured by the boll-weevil and
the cotton root rot fungus (Phymototrichum omnivorum) than if
grown under moist conditions.

TorPoGRAPHIC FEATURES

Beginning at sea-level in the low grassland and swampy areas
of the coast, the Rio Grande Plain on the average rises at the
rate of approximately five feet per mile. The coast is protected
by Padre Island, a very narrow strip composed mostly of dune
sand, extending along the coast from the mouth of the Rio Grande
to Corpus Christi, and separated from the mainland by Laguna

Madre, which is four to ten miles wide and rather shallow. In

general the coastal area consists of beach with low dunes, beyond
which are barren salt flats and salt marshes. On the surface of
the delta there are numerous low mounds locally known as “clay
dunes” (21). In northern Willacy County the wind piles up true
dunes (some of them twenty-five feet high) and blows out de-
pressions, making the topography irregular. All of Cameron,
most of Willacy, and a small portion of Hidalgo counties are
included in the Rio Grande delta.

The greater part of Hidalgo and Starr counties is included in
the Hebbronville Plain. There is a low rise which includes the
towns of Mission, McAllen, San Juan, Pharr, and Donna, and a
limited area near Raymondsville. This is known as the Mission
Ridge. All of the Rio Grande Delta with the exception of this
ridge has been subject to flood in the past. The delta begins at
Pefiitas, and in ordinary floods the water breaks out here, cover-
ing the streets of the village and filling in all of the low places

south of the Mission Ridge. Most of the sand belt occurs north

45

VEGETATION OF RIO GRANDE VALLEY

CLOVER

1937]

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46 MADRONO [Vol. 4

of the Lower Rio Grande area, but the southern end extends into
northern Hidalgo County and covers a small corner of Starr and
a portion of Willacy County (text fig. 1). The bed rock of this
region is covered by wind-blown sand. There are some migrat-
ing dunes; others have been stabilized by growth of grass or
shrubs.

The Hebbronville Plain extends from the sand belt and the
Rio Grande Delta west and north to a rather spectacular topo-
graphic feature known as the Bordas Scarp. The west face of
this escarpment averages sixty or seventy feet in height and is
composed of Oakville Sandstone and Frio Clay (text fig. 1).
The absence of erosion on the east side is probably accounted for
by the type of soil, which is a porous caliche overlain with loose
sand (35). The water leaches through so rapidly that little or
no erosion takes place. Looking at the Bordas Scarp from the
west, it is not unlike some of the flat-topped buttes of New
Mexico. More than half of Zapata County is included in the
Aguilares Plain (35), which is sixty or seventy feet lower than
the west side of the Hebbronville Plain and largely covered with
grass and mesquite.

The erosional valley of the Rio Grande begins at Pefiitas and
extends through Zapata County. Numerous arroyos drain this
part of the area, and the rough, broken region where they join
the river is known as the “Breaks of the Rio Grande.” This belt
varies in width from one to fifteen miles, averaging about seven.
In a portion of its course the river lies on an older valley filled
with detritus in which terraces have been cut. At other points,
particularly near Zapata and San Ignacio, the river runs along
high cliffs and bluffs of the Reynosa formation.

La Sal Vieja and Sal del Rey (text fig. 2) are salt lagoons.
It is thought (35) that the salt has been blown inland from the
Gulf.

GEOLOGY

The oldest formations are the Cook Mountain and the Cock-
field formations (35). They belong to the Claibourne group in
the Eocene and extend beyond the limits of Zapata County to the
north. The Fayette Sandstone, probably Oligocene, outcrops in
western Starr County and covers about one-half of Zapata County,
conforming somewhat in extent to the Aguilares Plain. A rather
narrow, irregular area of Frio Clay, with an outcropping of Oak-
ville Sandstone near Rio Grande City, overlies the Reynosa for-
mation forming the Bordas Scarp. The Oakville belongs to the
Miocene, and the Frio Clay is probably Oligocene.

The Reynosa formation covers the greater part of Starr
County and extends along western Hidalgo County. There are
small outcroppings elsewhere (text fig. 1). This formation com-
prises a greater part of the Hebbronville Plain. The Reynosa
is probably Pliocene and is composed of an indurated gravel

47

VEGETATION OF RIO GRANDE VALLEY

CLOVER

1937]

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48 MADRONO [Vol. 4

cemented by limestone, limestone with embedded pebbles, sand,
gravel, and sandstone. Lissie gravel occupies the central part of
Hidalgo County. It is of Pleistocene age and is composed chiefly
of unconsolidated gravel, balls of clay or irregular masses, lenses
and beds of sand and thin beds of limestone. Gravel from pits

near Sam Fordyce and Havana has been used for road-making ~

throughout the Valley.

The area of wind-blown sand and beach deposits is a wide
stretch reaching beyond Falfurrias and appearing in the Lower
Rio Grande Valley. The Prairie-Grass region and this area
almost coincide. Beaumont Clay of the Pleistocene begins at
Pefiitas, lying east of the Lissie Gravel region and joining the
fluviatile deposits which have their origin near Rio Grande City
as a narrow strip along the river. These deposits widen out to
many miles toward the coast.

DRAINAGE

With the exception of the Arroyo Colorado, the Rio Grande
is the only permanent stream. The upper part of the Lower Rio
Grande Valley is drained by arroyos (text fig. 2), in which there
are streams only following infrequent downpours, the deep,
ragged cuts testifying to the force of the torrents. Since these
arroyos are not bridged, traffic is sometimes forced to wait for
hours for the water to go down.

The poor drainage near the coast has been improved within
recent years by the construction of large, open drainage ditches.
Poor drainage has been a problem of great concern because much

of the irrigated land with insufficient drainage became so im- —

pregnated with alkali salts that some crops could no longer be
grown. New flood channels and levees, better canal systems and
more efficient handling of irrigation water have improved this
condition appreciably.

The Arroyo Colorado, a distributary of the Rio Grande, is a
deeply cut flood channel starting near Mercedes in eastern Hi-
dalgo County, and continuing in a northeasterly direction through
Cameron County to Laguna Madre. The water is salty as far
inland as Harlingen because the bottom of the channel is below
sea-level and there is little flow (3). Nearly all of the run-off
water of Cameron County flows into the estuaries of Laguna
Madre. )

In part of Zapata County there is no valley on the Texas side
of the Rio Grande, the river flowing against high cliffs (pl. IX).
In western Hidalgo County the valley has attained the width of
perhaps a mile. Lower down it becomes eight miles wide. In
western Cameron County the delta is many miles wide, and
excess water moves in poorly defined channels across the almost
flat surface toward the coastal lowlands. Some of these are
resacas and esteros, low places formed by the old river channels.

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 49

The writer’s survey of the vegetation of the Lower Rio Grande
Valley is based upon field studies made at three different periods
and extending over most of the four seasons. The periods of
study were as follows: June 30, 1932, to September 15, 1932;
February 25, 1932, to July 8, 1933; November 30, 1933, to Feb-
ruary 18, 1934. Information necessary for making a vegeta-
tional map was obtained by traverses on roads and trails. In
general, the roads are fairly good, but some transportation difh-
culties were encountered, especially in Zapata County. The
seasons of the study happened to be very dry; consequently,
many herbaceous plants which normally occur during favorable
seasons were missing. It was found that in this area vegetation
is influenced markedly by edaphic factors, and that it conforms,
more or less, to geological formations (text figs. 1, 3).

Locat Ecotocicat NOMENCLATURE

The Spanish-speaking people of Mexican stock in southern
Texas have developed a very definite and satisfactory classifica-
tion of the vegetation. Professor Bartlett (ms.) introduced
local nomenclature into the description of vegetational types in
his account of the botany of the San Carlos Mountains of Tamau-
lipas. He (2) and Lundell (27) have adopted this system in a
phytogeographical study of the Yucatan Peninsula. In the pres-
ent article local Spanish nomenclature is used to designate phyto-
geographic divisions, associations, and societies within the asso-
ciation. The writer has spent many years along the Rio Grande
and has learned these names directly from Texans and Mexicans.

Frequently the association or society is named for the domi-
nant species. For instance, mesquital applies to the association
dominated by the mesquite (Prosopis juliflora var. glandulosa) .?
A society within this association in which the mesquite is domi-
nant is also called mesquital. Zacatal derived from gzacate
(“grass”) designates the grassland association. This name is
used particularly for prairie grass. The grassland area is known
as los llanos. The coastal marsh grass (Spartina Spartinae) is
known as sacahuista, so the community which this species domi-
nates is a sacahuistal, and the area is called llano salitroso. That
part of the salt marsh occupied by water, and in which aquatic
plants occur is designated as badilla salitrosa, and sometimes as
pantafio. A charco is a depression or low place which supports
a growth of cat-tails (Typha latifolia), charas, sedges, and other
aquatics; while the resaca is a larger, more permanent body of
water formed by the old river channel. The borders of the
resacas are sometimes dominated by Parkinsonia aculeata (retama),
and the plant group is called a retamal. Other resaca communities

2 Since a systematic list of the species referred to forms a part of this
paper, space is saved in the ecological discussion by omitting the authorities
for scientific names.

50 MADRONO [Vol. 4

are named huisachal for Acacia Farnesiana (huisache); or mimosal
for Mimosa strigillosa. One resaca in Cameron County near Rio
Hondo has Ipomea fistulosa (amor) growing abundantly with
both Parkinsonia aculeata and Acacia Farnesiana. This is a rather
unusual situation, since Ipomea fistulosa is an escape and even
though the species is equalled in dominance by the other two, the
plant group receives the name amoral.

Hilly territory, chiefly the Bordas Scarp region, is known as
lomeria; and the high dry land in the upper counties and away
from the river is the mesa. The lomeria and parts of the mesa
have a characteristic growth consisting mostly of spiny shrubs
and stunted trees including many species. The Mexicans call
this growth chaparral, named from Acacia amentacea or chaparro
prieto (black chaparral) which is usually an important part of
the vegetation. A small area, usually a limestone or gravel hill,
dominated by this species is also a chaparral.

This use of the term chaparral differs from that of Cooper
(12) who refers to broad-sclerophyll scrub. He explains that
the word ‘chaparral’ is of Spanish origin meaning oak scrub.
In California and parts of Mexico this is generally true, but it
is not in south Texas. Chaparro prieto or black chaparral is the
Spanish name for Acacia amentacea, and any phytogeographic
division large or small which this species dominates is referred
to as chaparral. ‘This term is also used to designate an area cov-
ered by a scrubby growth in which the mesquite is inconspicuous
or absent. Authors usually include the mesquite in a chaparral
especially in localities in which the species is stunted and
serubby. Since this paper is following the local Spanish nomen-
clature for these ecological areas it is necessary to exclude the
mesquite. The author feels justified in using the name chaparral
for the semi-desert brush in view of the fact that it has been
used in this sense by several ecologists and other botanists for
many years. Emory (17) refers to the vegetation along the
Rio Grande as chaparral. Warming (36), Drude (16), Bray
(5), Engler (20), Harshberger (23), and Tharp (38), are other
authors giving this interpretation to the word.

Palma is the Spanish name for palm, and the palm grove or
small forest formation near Brownsville is known as boscaje de
palma.

In general the suffix -al, added to the Spanish name of the
plant in dominance (sometimes with vowel elision) designates
the plant group. An exception to this is nopalera, the name used
for an association and also for a society of Opuntia (nopal).

Such nomenclature as described will have no value in regions
where the Spanish language is not spoken; but in Spanish-
speaking localities an acquaintance with this method is impor-
tant. Bartlett (2) considers that much of this folk knowledge
can be systematically formulated in an ecological study. It is

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 51

Ficure 1

Figure 2

Pratt IX. Fig. 1. Near San Ignacio in Zapata County the Rio Grande
flows along steep bluffs. This hilly area broken by arroyos is known as the
“Breaks of the Rio Grande.” Fig. 2. Yucca tenuistyla (in the foreground)

growing in open brush-land. This plant is abundant in the vicinity of San
Ignacio.

52 MADRONO [Vol. 4

my purpose to use it in connection with terms ordinarily applied
to vegetational groups.

The writer is intentionally avoiding the use of many eco-
logical terms in English since they are often confusing even
when not accompanied by a local terminology. The association
is used in this paper to designate the larger, distinct plant
groups, and society for smaller groups within the association.
The word community is used for groups in which the rank
is uncertain (37).

PHYTOGEOGRAPIC STUDY

There is no true tropical area in Texas. The freezing tem-
peratures which sometimes occur limit genuine tropical life to a
few insects, reptiles, and birds (1). However, near the mouth
of the Rio Grande and extending inland some distance are a few
plants of generally tropical distribution. One would place Sabal
texana foremost in this list. Others are Daubentonia longifolia,
Lantana horrida, Malphigia glabra, Amyris parvifolia, Helietta parvi-
folia, Schaefferia cuneifolia, Serjania incisa, Cardiospermum Halica-
cabum, Ruellia tuberculosa, and Avicennia nitida.

The Rio Grande Valley is of much interest botanically be-
cause plants representing western desert, northern, coastal, and
tropical floras are all found in a relatively small area.

I. Tue Mesguitat Criimmax

Prosopis juliflora var. glandulosa is classed as a chaparral plant
in parts of the United States where it is shrubby. In the Rio
Grande Valley it often reaches a height of thirty-five feet and is
sometimes two feet or more in diameter. Here the mesquite is
a tree, and any association or plant group in which the mesquite
is dominant or especially conspicuous is known as mesquital.
The mesquital covers most of the area and exhibits several
phases (text fig. 3). }

Mesguirat-zacaTaL. This phase covers the greater part of
Zapata County, conforming rather closely to the area known as
the Aguilares Plain. It is interrupted by the Bordas Scarp, but
continues on the other side as a narrow strip through the north-
ern part of Starr County and across Cameron County to the
Gulf.

Most of this territory is a flat sandy plain. Even though the
area west of the Bordas Scarp has the same dominating eco-
logical characters as that on the east side, the composition of the
vegetation is quite different. The dominants in this western
part in Zapata County are: Prosopis juliflora var. glandulosa, with
Bouteloua barbata and Aristida purpurea probably dominant among
the grasses, although it is difficult to name dominants where over-
grazing interferes. Other grasses enter this phase of the
mesquital and there are many societies of mixed shrubs occurring
with varying dominance. The most important dominants are:

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 53

Zizyphus obtusifolia, Leucophyllum frutescens, Acacia Berlandiert,
Bumelia lycioides, Celtis pallida, Schaefferia cuneifolia, Forestiera
angustifolia, Acacia amentacea, and Cercidium texanum. ‘These
shrubs are often scattered through the open mesquite woods and
occasionally form a heavy growth near an arroyo.

Yucca tenuistyla is rather abundant northeast of San Ignacio
forming societies in open grassy areas. Such a society is known
as an aguapal. This species is often found in open brush-
land (pl. IX).

Herbaceous plants form many of the societies. The most im-
portant of these are: Gaillardia pulchella, Lupinus texensis, Oeno-
thera laciniata var. mexicana, Parthenium Hysterophorus, Verbesina
encelioides, Aphanostephus skirrobasis var. Halli, and Jatropha
spathulata.

By far the greatest portion of the mesquital-zacatal lies east of
the Bordas Scarp. The dominants here are: Prosopis juliflora var.
glandulosa and probably Aristida purpurea, Eragrostis curtipedicel-
_ lata, and Eragrostis secundifiora. It is difficult to name a single grass
which dominates the entire phase, for there is varying dominance
depending upon edaphic factors and light conditions. In poorer
soil the grama grasses (Bouteloua) grow with little competition ;
in other situations they form the layer under taller grasses such
as Eragrostis curtipedicellata, Eragrostis secundiflora, Chloris cucul-
lata, and Aristida purpurea.

Other herbaceous plants forming societies are: Callirrhoé digi-
tata, Commelina crispa, and Jatropha stimulosa. Shrubs invading
this phase are: Celtis pallida, Zizyphus obtusifolia, Leucophyllum
frutescens, Acacia amentacea. Societies and clans of Opuntia Lind-
heimeri and Opuntia leptocaulis are occasionally found.

Echinocereus angusticeps, recently described (10) as a species
distinct from EF. papillosus, occurs near Linn, and is found no-
where except in this limited area. It would be interesting to
know the conditions which make this locality especially favorable
for the growth of this species. The only other cacti found here
are: Dolicothele sphaerica, Neomammillaria hemisphaerica, Opuntia
Lindheimeri, and O. leptocaulis. Plants other than cacti are: Con-
volvulus incanus, Jatropha spathulata, Prosopis juliflora var. glandu-
losa, Zizyphus obtusifolia, Celtis pallida, Aristida purpurea, and
Bouteloua barbata.

The mesquite trees are often infested with mistletoe, Phora-
dendron flavescens. In the region of Zapata and San Ignacio
many of them have been seriously damaged by this parasite (6).
Around Aguilares the infestation is particularly heavy, and many
trees are dying.

MESQUITAL-NOPALERA. It is impossible to draw a definite line
between the phases of the mesquital area. An attempt has been
made (text fig. 3) to show the merging of one phase into the
other. There is a narrow strip from east to west in which condi-
tions seem optimum for the growth of opuntias. They vie with

5A MADRONO [Vol. 4

mesquite for dominance, and, in some places, crowd out most ©
other species. Celtis pallida, Viguiera stenoloba, Lippia ligustrina,
Lantana horrida, and Bouteloua barbata are commonly found even
in the densest growths. Cleared land which has been abandoned
is soon covered by a heavy growth of Opuntia. The opuntias
readily reproduce vegetatively, a single joint being capable of
giving rise to one or more new plants. Since these joints are
rather easily broken from the parent plants the spread of the
genus in a free area is extremely rapid. The soil is a deep sandy
loam. There is a gentle slope to the east, providing good drain-
age, and here the mesquite is probably more abundant than in
any other part of the Rio Grande Valley.

Included in the mesquital-nopalera there are two depressions
known as Sal del Rey (Hidalgo County) and La Sal Vieja (Wil-
lacy County). The sparse vegetation at the margin of the lake
is of the beach and coastal prairie type. Monanthochloé littoralis,
Strombocarpa cinerescens, and Chenopodium album are most abun-
dant.

MeEsQuiTAL-cHAPARRAL. The invasion of the chaparral vegeta-
tion is comparatively recent and thought to be the result of over-
grazing and drought. The mesquital-chaparral is by far the most
important phase of the mesquital. It covers the greater part of
the territory in the Lower Rio Grande Valley, and in general typi-
fies the vegetation which is found in much of southwestern Texas.

There is a marked variation in soil conditions here, and this
difference affects the size and luxuriance of the growth. Many
species occur throughout the range, and others are limited in
extent.

On the surface of the delta in eastern Cameron County there
are numerous small mounds locally known as “clay dunes.”
They project several feet above the coastal plain and are often
covered with a dense growth of chaparral and mesquite (pl. X).
This growth is similar in structure to much of the vegetation
between Brownsville and San Benito, but the shrubs near the
coast are characteristically twisted as a result of heavy winds.
The association has no general dominant. Some definite societies
within it are dominated respectively by the following species:
Siderocarpos flexicaulis, Leucophyllum frutescens, Zizyphus obtusi-
folia, Castela Nicholsonii, Randia aculeata, Forestiera angustifolia,
Prosopis juliflora var. glandulosa, and Celtis pallida. <A high ridge
near the Rio Grande supports a much sparser growth of shrubs
interspersed with mesquite and the following cacti: Ferocactus
hamatacanthus, Echinocereus pentalophus, Ancistrocactus Scheeri,
Hamatocactus setispinus, and Neomammillaria hemisphaerica.

These “islands” of chaparral are separated from the main
chaparral-mesquital area by the sacahuistal and a transition zone.
There is at first an invasion of mesquite into the coastal prairie
area. Lycium carolinianum, Celtis pallida, and Zizyphus obtusifolia
are usually the first shrubs to enter. The composition of the

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 55

Figure 2

Pratt X. Fig. 1. Chaparral vegetation covering a “clay dune” near the
coast. The dominant shrub with dark foliage is Siderocarpos flexicaulis. (See
also Pl. XIII, fig. 2.) Fig. 2. The chaparral-cenizal near Roma, Starr County.

The light areas are dominated by Leucophyllum frutescens; the dark areas
are dominated by Acacia amentacea.

56 MADRONO [Vol. 4

mesquital-chaparral farther inland is similar in parts of Hidalgo
and Willacy counties and in Cameron County. The brush is
usually five to eight feet tall and rather dense. Dominance is
dependent on several conditions, and is difficult to determine in
much of this area. The flora is more or less characteristic
throughout, but it is varied in its distribution and aspects by sev-
eral factors.

Soil and drainage have a decided effect upon the distribution
of plants in this association.

Limestone outcrops occur as hills surrounded by the Reynosa
formation. Leucophyllum frutescens and Acacia amentacea are
dominant on these hills in a society known as chaparral-cenizal
(pl. X). Other prominent plants are Bouteloua trifida, Panicum
Hallu, Jatropha spathulata, Prosopis juliflora var. glandulosa, Kar-
winskia Humboldtiana, Hamatocactus setispinus, Neomammillaria
hemisphaerica, and Opuntia Lindheimeri.

Three species, Leucophyllum frutescens, Acacia amentacea, and
Acacia Berlandiert share dominance on limestone and Reynosa
gravel near the Rucio ranch, northeast of Rio Grande City. Other
plants occurring in abundance are Aristida purpurea, Bouteloua bar-
bata, Condalia obovata, Karwinskia Humboldtiana, Bumelia angusti-
folia, Echinocereus papillosus, Opuntia Lindheimeri, and Aphanoste-
phus skirrobasis var. Hallit.

Mortonia Greggu (pl. XI) is dominant in societies on hillsides
in a limited area five miles north of La Joya. <A few plants are
also found near Lake La Joya. The outcrop in the La Joya
vicinity is Lissie Gravel. A few stunted plants of the above
named species also occur in a small outcrop of Lissie Gravel near
Rio Grande City.

A sandstone and gravel outcrop on a hill south of Mission,
known as La Lomita, shows no dominant. There is a dense
growth of mixed vegetation which contains a greater variety of
plants than is usually found in an equal area in the mesquital-
chaparral. The following plants are prominent or abundant in
this society: Adelia Vaseyi, Bernardia myricaefolia, Schaefferia
cuneifolia, Coursetia axillaris, Acacia amentacea, Prosopis juliflora
var. glandulosa, Randia aculeata. Coursetia azillaris is extremely
rare, probably occurring nowhere else in the Rio Grande Valley.

The amargosal is an area dominated by Castela Nicholsonit.
This species is particularly abundant in Lissie Gravel near Pefiitas
and La Joya (pl. XI). Sub-dominants are Acacia amentacea and
Zizyphus obtusifolia. Other plants of importance are Celtis pal-
lida, Karwinskia Humboldtiana, Schaefferia cuneifolia, Ancistrocactus
Scheeri, Dolicothele sphaerica, Echinocereus enneacanthus, E. penta-
lophus, Hamatocactus setispinus, Homalocephala texensis, Neomam-
millaria hemisphaerica, N. Heyderi, Opuntia leptocaulis, O. Lind-
heimeri, O. Schottii, Guaiacum sanctum, and Viguiera stenoloba.

High dry land included mostly in the Hebbronville Plain is
known as the mesa. The soil is sandy loam with limestone and

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 57

Figure 1

Figure 2

Puate XI. Fig. 1. A limestone outcrop. Mortonia Greggii (foreground)
dominant, with Leucophyllum frutescens, Cordia Boissieri and Acacia Ber-
landiert present. Fig. 2. Lake La Joya bordered by a dense growth of
Castela Nicholsonti. This society is known as amargosal.

58 MADRONO [Vol. 4

gravel outcrops. Much of the vegetation is mixed, with no out-
standing dominant. The ebanal is dominated by Siderocarpos
flexicaulis; the nacahuital by Cordia Boissiert and the comal by
Bumelia lycioides. Cordia Boissiert is usually found to prefer
limestone and gravel hillsides. Lippia ligustrina and Celtis pal-
lida are usually prominent here. Bumelia lycioides usually occurs
on high level land in sandy loam soil, although single specimens
are scattered throughout the Rio Grande Valley.

Prosopis juliflora var. glandulosa often dominates societies
throughout the mesa, especially in sandy soil or in lower places
between hills or near arroyos. In this mesquital the usually
prominent species are Celtis pallida, Opuntia leptocaulis, and Opun- |
tia Lindheimeri. The latter likewise attains dominance in certain
localities, often almost excluding other species.

East of the mesa and in the western part of the Rio Grande
Delta there is a variety of societies largely determined by soil
and moisture conditions. Much of the vegetation in this area is
mixed, with no outstanding dominant. Prominent plants grow-
ing in the Harlingen Clay near San Benito are given below
grouped according to layers: (1) lower layer, Setaria macro-
stachya, Trichloris pluriflora, Croton Cortesianus, Opuntia Lindhei-
meri, Salvia coccinea, Parthenium Hysterophorus, and Viguiera steno-
loba; (2) upper layer (shrubs 5-8 feet high, mesquites 10-15
feet high) Celtis pallida, Forestiera angustifolia, Leucophyllum fru-
tescens, Pithecolobium brevifolium, Prosopis juliflora var. glandulosa,
Bumelia angustifolia, Zizyphus obtusifolia, Opuntia Lindheimeri, Lan-
_ tana horrida, and Heimia salicifolia.

Acanthocereus pentagonus (pl. XII), commonly called “night-
blooming cereus’’, with the mesquite forms an association within
the mesquital-chaparral in the vicinity of Rio Hondo. This asso-
ciation is several miles in extent. About four miles east of Rio
Hondo the cactus is dominant, climbing in mesquites and form-
ing such a dense tangle that it is almost impossible to get through
without a “machete.” Farther north, in an open mesquite wood
it is frequent with Spartina Spartinae and Opuntia Lindheimeri. In
places where the cactus and mesquite share dominance the fol-
lowing are prominent in the upper layer: Celtis pallida, Zizyphus
obtusifolia, Xanthorylum Pterota and Opuntia Lindheimeri. The
lower layer is composed largely of the following: Rivina humilis,
Malvastrum caromandelianum, Sida paniculata, Heliotropium indicum,
Gilia incisa, Hamatocactus setispinus and Salvia coccinea.

Depressions anywhere in the lower part of the Rio Grande
Valley in which run-off and flood waters gather often. have a
growth of Parkinsonia aculeata (retama). A society dominated by
this species is called retamal. Acacia Farnesiana is often second in
dominance here, and in depressions which are periodically dry it
is apt to be dominant. Typha latifolia, Scirpus Halli, Atamosco
terana and Castalia elegans (pl. XII) are usually present growing

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 59

Figure 2

Pirate XII. Fig. 1. Acanthocereus pentagonus. This cactus requires
much more water than most species. It extends over an area of several square
miles east of Rio Hondo and often stands in water for weeks after a gulf
storm without serious injury. Fig. 2. A shallow resaca near Alamo. Scirpus
Halli and Castalia elegans are shown here. Parkinsonia aculeata is in the
background.

60 MADRONO [Vol. 4

in the water. Some societies also contain Echinodorus cordifolius
and Sagittaria variabilis.

The huisachal is dominated by Acacia Farnesiana. Besides the
ones mentioned above there are other societies dominated by this
species near the Rio Grande. It survives flood water, in which it
stands for months, and the soil when drying out often develops
deep cracks making conditions difficult for the growth of small
plants. Rumex meaxicanus, Hartmannia speciosa, Ambrosia elatior,
Parthenium Hysterophorus and Aster exilis are among the first to
enter.

The charco is a small low area covered with shallow water.
Typha latifolia is dominant associated with Cyperus articulatus,
Scirpus validus, Marsilia vestita, Naias guadalupensis, Hartmannia
speciosa and Aster eailis.

Resacas formed by cut-offs in the old river channel usually
contain shallow water. These resacas have practically the same
type of vegetation which is found in other depressions. A resaca
near Rio Hondo has an abundance of Ipomea fistulosa (probably
an escape) occurring with the usual dominants. This society is
called amoral. Aquatics here are Chara praelonga, Marsilia vestita,
M. macropoda, Typha latifolia, Lemna minor, Heteranthera limosa,
Eichornia crassipes, Castalia elegans, Utricularia subulata, Radicula
Walteri and sedges (named at end of paper). Wet bank vegeta-
tion includes various sedges, Cynodon Dactylon, and Aster exilis.
Lepidium virginicum, Hartmannia speciosa, Urtica chamaedryoides
and Lycopersicon cerasiforme occur in the moist shady zone.

Irrigation has had some effect on plant distribution. Seeds
are carried by the water and become established along the banks.
The moisture added to the soil permits the growth of plants
ordinarily excluded by dry conditions. The main canals are
always filled with water and such species as Cynodon Dactylon,
Holcus halepensis, Paspalum Langei, Commelina longicaulis and Par-
thenium Hysterophorus grow along the banks. Artificial depres-
sions, locally called “‘barrow pits’’ at the sides of the canals, al-
ways contain some seepage water, and usually have the follow-
ing species: Marsilia vestita, Typha latifolia, Echinodorus cordi-
folius, Phragmites communis, Cyperus acuminatus, Cyperus oxycario-
ides, Scirpus validus, Castalia elegans, and Jussiaea diffusa. Some of
the smaller canals which carry water only periodically are
choked with FHichornia crassipes. Typha latifolia, Salix longifolia,
Celtis pallida, and Cynodon Dactylon also flourish in this environ-
ment.

There is a heavily wooded area along the Rio Grande in
Cameron County in which trees reach a height of fifty feet or
more. Near Alamo in such a situation the upper layer consists
of Celtis mississippiensis, Ulmus crassifolia, Siderocarpos flexicaulis
(pl. XIII), Sapindus Drummondi (pl. XIII) and Frazinus Ber-
landieri. These trees are festooned with Tullandsia usneoides.
Tillandsia recurvata and Tillandsia Baileyi are also present. Promi-

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 61

Ficure 1

Figure 2

Prare XIII. Fig. 1. Wooded area near the Rio Grande, Alamo. The
large tree is Sapindus Drummondii. Fig. 2. Siderocarpos flexicaulis in woods
near the Rio Grande, Alamo. This same species is a small shrub in dry areas.

62 MADRONO [Vol. 4

nent species in the middle layer are Celtis pallida, Zizyphus obtusi-
folia, Porlieria angustifolia, Malphigia glabra, Diospyros texana and
Bumelia angustifolia. The lower layer is a tangle of vines and
weak-stemmed, herbaceous plants. The most prominent ones
are Rivina humilis, Clematis Drummondu, Urtica chamaedryoides,
Cuscuta indecora, Cardiospermum Halicacabum, Plumbago scandens,
Capsicum baccatum and Monarda dispersa (pl. XIV).

Flood waters disturb and destroy the smaller vegetation along
the river leaving stretches of deep sand. These stretches are soon
covered by almost pure stands of Baccharis which are often swept
away before being replaced by other species.

A single specimen of Taxodium mucronatum stands in the
woods south of Havana. The tree is approximately forty feet
tall. Mexican residents report that it has been there for at least
a hundred years. Emory (17) states that cypress was found
near the mouth of the Rio Salado, which is a tributary of the Rio
Grande, and expressed the hope that some future day this species
would spread down the river, furnishing building material for
inhabitants.

II. Tue CHaprparraLt CLIMAx

The Bordas Scarp and a strip along the Rio Grande between
Roma and San Ignacio is covered with a low shrubby growth
which seems fairly stable. There are societies of mesquite on
flats and along arroyos, but in general the hilly broken territory
known as the “Breaks of the Rio Grande” is a distinct chaparral,
as the term is used in this paper. Acacia amentacea is probably a
general dominant with subdominants varying with the locality.
Some of the most outstanding associations, societies and com-
munities are given below.

CrenizaL. This association is dominated by Leucophyllum fru-
tescens with subdominants Acacia amentacea and Acacia Beérlandiert.
The presence of Leucophyllum usually indicates a limestone out-
crop, and Acacia Berlandieri is often found on gravel hills. A
cenizal occurs about two miles west of Roma and is several miles
in extent (pl. X). The following plants were collected in this
area: Ephedra antisiphilitica, Atriplex acanthocarpa, Bouteloua bar-
bata, B. trifida, Cenchrus pauciflorus, Pappophorum mucronulatum,
Talinum angustissimum, Acacia Berlandieri, A. amentacea, Parosela
nana, Croton ciliato-glandulosus, C. fruticulosus, Jatropha spathulata,
Schaefferia cuneifolia, Karwinskia Humboldtiana, Microrhamnus eri-
coides, Abutilon incanum, Gayoides crispum, Sida filipes, Ancistro-
cactus Scheeri, Hamatocactus setispinus, Echinocereus enneacanthus,
Neomammillaria hemisphaerica, Opuntia leptocaulis, O. Lindheimeri,
Heliotropium confertifolium, Marilaunidium hispidum, Goniostachyum
citrosum, Nicotiana repanda, Leucophyllum frutescens, Actinea
odorata.

Baretta. This occurs on gravel and limestone hills. The
dominant species is Helietta parvifolia (pl. XIV). It is a rather

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 63

Ficure 1

Figure 2

Prats XIV. Fig. 1. A society of Monarda dispersa in a mesquite wood,
Havana. Fig. 2. Helietta parvifolia on a low hill east of Rio Grande City.

64 MADRONO [Vol. 4

large community extending over several hills east of Rio Grande
City. A few scattering plants are found as far east as Pefiitas,
and some grow in the region of the petrified forest east of Roma.
Other prominent plants in this community are as follows: Boute-
loua barbata, Setaria macrostachya, Agave Lechuguilla, Acacia amen-
tacea, Cercidium floridum, Castela Nicholsonu, Koeberlinia spinosa,
Ancistrocactus Scheeri, Coryphantha Runyonii, Opuntia leptocaulis, O.
Lindheimeri, Thelocactus bicolor, Coldenia canescens, Cordia Boissieri,
Lippia ligustrina. Agave Lechuguilla is not abundant but is worthy
of mention as this locality seems to be the eastern limit of its
range. A new species, Coryphantha Pirtleana, was recently
found here by Pirtle brothers of Alamo.

Hecutia Texensis Society. The soil may be somewhat saline
since Varilla texana, which is a halophyte (14), is always present.
Burros enjoy eating the succulent flower stalks of Hechtia, many
of which are prevented from seeding by being nipped off early.
New plants are produced at the base of the parent plant, forming
large mounds so dense that no other species can gain a foothold
(pl. XV). Some of the plants besides Varilla that are commonly
found with Hechtia are Panicum Halli, Zizyphus obtusifolia, Sida
filipes, Schaefferia cuneifolia, Jatropha spathulata, Porlieria angusti-
folia, Pappophorum bicolor, Echinocereus enneacanthus, and Opuntia
Lindheimeri.

Agave Society. This society occurs on sandy loam and lime-
stone hills. Agaves are found in only a few localities in the two
upper counties. Probably the largest group is five miles south of
Zapata. The vegetation growing with this species is character-
istic of that found in much of the chaparral (pl. XV).

Borpas Scarp Vecetation. The approach to the Bordas Scarp
is a gentle slope toward the west. The vegetation is of the mes-
quital-chaparral type, changing to the lower, sparse chaparral found
on gravel. An area about a mile in length was chosen as repre-
sentative of the chaparral of the escarpment. An interesting
cactus, Astrophytum asterias, is found here, which, as far as
known, occurs nowhere else in the United States (9). The fol-
lowing species were collected in this locality: Ephedra antisiphi-
litica, Aristida purpurea, Chloris cucullata, Eragrostis curtipedicellata,
Yucca sp., Runyonia longiflora, Celtis pallida, Talinopsis frutescens,
Acacia amentacea, Prosopis juliflora var. glandulosa, Jatropha spathu-
lata, Schaefferia cuneifolia, Karwinskia Humboldtiana, Zizyphus ob-
tusifolia, Koeberlinia spinosa, Menodora heterophylla, Astrophytum
asterias, Coryphantha Runyonii, Dolicothele sphaerica, Echinocereus
enneacanthus, E. Fitchu, E. pentalophus, Hamatocactus setispinus,
Lophophora Williamsit (pl. XVI), Neomammillaria hemisphaerica, N.
Heyderi, Opuntia leptocaulis, O. Lindheimeri, Thelocactus bicolor,
Wilcoxia Poselgeri, Lippia macrostachya, Ibervillea Lindheimeri, Va-
rilla texana, Verbesina encelioides, Zinnia pumila. The following
plants were found at the base of the escarpment: T'riodia mutica,

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 65

Ficure 1

Figure 2

Prats XV. Fig. 1. A society of Hechtia tewensis near an arroyo be-
tween Roma and Zapata. Fig. 2. Agave melliflua in flower. This society
occurs in a mixed chaparral near Zapata.

66 MADRONO [Vol. 4

Sporobolus Wrightii, Rivina humilis, Jatropha Berlandieri, Abutilon
Wright, Amoreuxia Wright, Aphanostephus skirrobasis var. Halli,
Dyssodia tephroleuca, Simsia calva, Zexemenia hispida.

Costittat. The distribution of this society is mostly in Za-
pata County. The dominant is Microrhamnus ericoides. In places
this species is so dense as to almost crowd out competitors. Low
mesquites, Zizyphus obtusifolia and Colubrina texensis are fre-
quently associated with it.

Pato VerRpE Society. The dominant species are Cercidium
texanum in the region of Zapata, and Cercidium floridum near
Pefiitas. These species usually occur on dry mesas in open brush-
land. They thrive in dry gravel soil as the leaves are early de-
ciduous and the stems carry on photosynthesis. Bouteloua bar-
bata, B. trifida, Coldenia canescens, Heliotropium confertifolium, Par-
thenium Hysterophorus are some of the herbaceous species com-
monly associated with Cercidium. The shrubs of this society vary
with the locality.

GoBERNADORAL. The dominant, Covillea tridentata, has in-
vaded northern Zapata County from the northwest. The fact
that it is most abundant on very dry and unproductive soil might
give the impression that it prefers that habitat, but it does not.
Covillea is easily crowded out by other vegetation in more favor-
able situations. Spaulding (32) states that it is capable of adapt-
ing itself to desert conditions by reducing leaf surface and by
abstracting water from very dry soil, but it is capable of living
and does live as an ordinary mesophyte if a suitable supply of
water is available.

VEGETATION ALONG ArRoyos. Shrubs inhabiting areas cut by
arroyos are often larger near the banks where water stands after
a storm. Species such as Acacia Farnesiana and Parkinsonia acule-
ata grow near these water-holes. Cynodon Dactylon, Pappo-
phorum bicolor, P. mucronulatum, Sporobolus argutus, Trichloris men-
docina, Triodia pilosa, Panicum Halli, Lepachys columnaris var.
pulcherrima, Nicotiana longiflora, Parosela nana, V erbesina encelioides,
and Chamaesyce laredana are plants growing in dry arroyo beds.
At Arroyo Loma Blanca Ferocactus hamatacanthus (pl. XVI) is
abundant. One plant has persisted in the crotch of a tree for at
least four years in spite of the fact that water has washed away
all the soil in which it originally grew. Echinocereus enneacanthus,
the species of cactus which has a greater distribution than any
other in the Lower Rio Grande excepting the opuntias, often
grows to the very edge of arroyos.

There are other societies in the chaparral which may also be
found in the mesquital-chaparral phase. Bumelia lycioides and B.
angustifolia occur in societies, clans and families throughout the
area. Such plant groups are known as comales (singular, comal).
Diospyros texana forms a gapotal, and Yucca a pital. There are
few areas in which Yucca grows abundantly. It is usually scat-
tered singly or in very small groups.

(To be concluded)

—

a tn

1937] CORY: TALINUM PULCHELLUM IN TEXAS 67

EARLY CALIFORNIA PLACE NAMES USED BY
DANIEL CLEVELAND

A copy of the following letter was received recently from Dr.
L. R. Abrams. Written in California thirty years ago by one of
the early botanical collectors, it gives important information con-
cerning certain old place names then in use. An account, by
Dr. W. L. Jepson, of the life and botanical work of Daniel Cleve-
land has been published in this journal (MaproNno 1: 267-268.
1929).—H. L. M.

San Diego, Cal., Feb. 7th, 1906
Mr. Le Roy Abrams,
U. S. National Herbarium, Washington, D. C.
My dear Sir:—

Your letter of the 2nd. inst. is just received. I take pleasure in
answering it. “Larken’s” (‘‘Pete Larkens,” it used to be called locally)
is, as you surmise, what is now known as Jacumba Hot Springs, be-
tween 75 and 80 miles east of this city and some half mile north of the
Mexican Boundary line. Peter Larkin, an American, owned and occu-
pied this cattle ranch from about 1870 to 1887. About the latter year
he removed to another ranch some 20 miles west of “Jacumba.” The
Jacumba Indian tribe lived in the immediate vicinity of “Larkins.”
The Jacumba Hot Spring is on the old Larkin ranch. Mountain
Spring is about 10 miles east of Larkins, or Jacumba and on the edge
of the desert. ‘“Tighe’s’”—so named from the owner of the ranch who
settled and lived there about 1875 and until about 1882—is situated
some six to eight miles east of the town of Ramona, which is nearly 40
miles from San Diego. This ranch was owned and occupied before
Tighe’s occupation by Billingsly, about 1873-5, and before 1873 by
Luckett, by whose names this ranch was also known, and is so named
by collectors. Dr. Edward Palmer and I used the name of “Luckett”
for this locality in connection with our plants collected in that vicinity.
The name of Larkin was used by Dr. Edward Palmer, by me and other
collectors for the country within 10 miles of Larkin’s ranch. Tally’s
ranch lies in a valley at the base of the South Cuyamaca Mountain
peak some 8 miles southward from the town of Julian in this county.
The name as used by collectors covers quite a large valley section sur-
rounding the old (James) Tally farm house... .

Sincerely yours,
Daniel Cleveland.

THE OCCURRENCE OF TALINUM PULCHELLUM
IN TEXAS
V. L. Cory

On a field trip to the southwestern part of the Glass Moun-
tains, northwest of Marathon, Brewster County, Texas, April 11,
1936, a plant with a solitary, purplish-red, showy flower, borne

68 MADRONO [Vol. 4

just above the surface of the ground, was seen for the first time
by the writer. Six of these plants were taken for observation
under cultivation, of which three were grown by H. B. Parks at
San Antonio and the other three by the writer at Sonora. These
grew readily, and proved to be species of Talinum. The solitary,
showy flowers with a spread of as much as three centimeters indi-
cated that the species was T’.. pulchellum Woot. & Standl. (Contr.
Was. Nat. Herbs 16.121. 19 03)).

Referring to the original description of this species and com-
paring the plants in our care, which were growing under favora-
ble conditions, discrepancies were noted: the peduncle was 15
millimeters long or, together with the pedicel, 30 millimeters long;
for the most part also, the peduncles were cymosely 3-flowered,
as contrasted with the original description in which the peduncles
were described as 2-8 millimeters long and only 1-flowered.
Furthermore, a count of stamens revealed 34 instead of about 20.
In the treatment of this species by Percy Wilson (N. Am. Fl. 21:
284-285. 19382) the number of stamens was corrected to read
25-38, but still it was stated that the flowers were solitary.

Transferring attention now to C. H. Mueller’s publication of
Talinum Youngae (‘““A New Species of Talinum from Trans-pecos
Texas,’ Torreya 33: 148-149. 19383) the description is seen to
fit our plant except in minor matters of measurement and in the
number of stamens, stated as being about 18. The separation
of the two species by C. H. Mueller (Torreya 34: 40-41. 1934)
is based upon the flowers being on longer peduncles and usually
3 in cymes in T'. Youngae, while in T. pulchellum the flowers are
solitary on pedicels about 1 centimeter long. A study of our
material transplanted from the Glass Mountains shows that these
differences do not hold constant. It is indicated therefore that
only one species is involved. The plants vary according to
the conditions under which they grow, longer peduncles and
3-flowered cymes tending to develop under favorable conditions.
It seems possible, also, that specimens taken late in the season
may show uniformly a reduced number of stamens.

In C. H. Mueller’s original publication (1933) the following
references are made to earlier collections of TI. Youngae by Dr.
M. S. Young: “Summit of Mt. Livermore, Davis Mountains,
August 15, 1914,” and “Very abundant on high, rocky slopes,
Pine Canyon, Guadalupe Mountains, August 15, 1916.” The
type locality of the original collection of T. pulchellum, by Dr. E. O.
Wooton, was also in the Guadalupe Mountains, “near Queen,
New Mexico, August 2, 1909.” It seemed desirable therefore, to
collect the plant on Mt. Livermore, and this was done on May 3,
1936. Several plants were taken on high rocky slopes. Each
had a solitary flower; a typical flower revealed 34 stamens, or
exactly the same as in the plant from the Glass Mountains.
There is no reasonable doubt that this plant is the same as that

1937] NOTES AND NEWS 69

formerly collected by Dr. Young. To the writer, there seems to
be sufficient evidence that both these collections should be re-
ferred to T. pulchellum Woot. & Standl. JT. Youngae C. H. Mueller
therefore becomes a synonym of T. pulchellum Woot. & Standl.

The following revised description of T. pulchellum includes
characters which develop under favorable conditions of growth.

TALINUM PULCHELLUM Woot. & Standl. Contr. U.S. Nat. Herb.
fo. 121. 19138. 7. Youngae C. H. Mueller, Torreya 33: 148.
19338.

Leaves up to 28 mm. long, 3 mm. in diameter, terete (not even
slightly flattened except under adverse conditions of growth),
blunt, densely and evenly distributed along the stem, but under
adverse conditions appearing basal; not narrowed at the base, at
point of attachment three-fourths to four-fifths the diameter of
the stem, subtended by a rounded, flattened margin, extending
around the base below the point of attachment; peduncles axil-
lary, stout, under favorable conditions 3-flowered, under other
conditions commonly 1-flowered, or sometimes 2-flowered; when
more than 1-flowered commonly with only one bloom open at one
time, flower opening about mid-afternoon and closing at sun-
down; when 3-flowered, peduncle up to 15 mm. long, at the apex
marked by a definite ring or sunken joint, which is subtended by
a pair of opposite bracts 5-6 mm. long, linear-lanceolate, more
or less fleshy, and scarious-margined, each bract subtending a
lateral pedicel; lateral pedicels equal in length to the middle or
terminal pedicel (15 mm. long), likewise terminating in a soli-
tary flower and bearing 5 mm. from the base a pair of opposite
bracts, similar to but shorter (4-5 mm. long) than the bracts sub-
tending the pedicels; middle pedicel ebracteolate.

The report that T. pulchellum is known only from the type
locality, which is near Queen, New Mexico, may now be modified
to show the distribution of this species throughout the mountains
of southwestern Texas.

Agricultural Experiment Station,
Sonora, Texas, August, 1936.

NOTES AND NEWS

Dr. Albert Levan of Lund University, Sweden, arrived in
Berkeley February, 19387. He will spend six months in Cali-
fornia continuing cytogenetical studies on the genus Allium. He
is at present engaged in field work in southern California and
while there, will make his headquarters at the California Insti-
tute of Technology, Pasadena.

At a meeting held in Santa Barbara on February 19, 1937,
plans were laid for the establishment of a branch of the Cali-
fornia Botanical Society in that city. Director Maunsell Van

70 MADRONO [Vol. 4

Rensselaer of the Blaksley Botanic Garden and Miss Ruth Hart-
well of the Santa Barbara Museum of Natural History were ap-
pointed as a committee to perfect the plans of organization.
The meeting was attended by forty-five enthusiastic botanists
and amateurs. Dr. Herbert L. Mason of the University of Cali-
fornia addressed the meeting on the subject of “The flora of the
Carpinteria asphalt deposits and its bearing on the history of
the Monterey forest.” The following Sunday a field trip was
made to Pine Canyon and Burton Mesa in northern Santa Bar-
bara County, a region little explored botanically. The follow-
ing persons attended the field trip: Dr. Herbert L. Mason; Pro-
fessor Woodbridge Metcalf; Mr. and Mrs. Sydney Anderson;
Mr. and Mrs. Charles C. Christiansen; Jan Christiansen; Meredith
Christiansen; Dr. Frances Long; Miss Louisa M. Long; Miss
Ella Mae Ottery; Miss Gil C. Pope; Mr. Emmett Martin; Mr.
E. D. Rowe; Miss Ruth Hartwell; Rev. Seraphin Muller; Mr.
and Mrs. Hugh Dearing; Mr. and Mrs. Maunsell Van Rens-

selaer; and Miss Patricia Van Rensselaer.

The following distributional notes have been contributed by
Mrs. Dorothy R. Harvey of San Diego State College. Agave
Shawi thought to be extinct along the coast near the Mexican
boundary was found on Point Loma, San Diego, about three
miles west of Ocean Beach. The colony extends about seventy-
five yards along the cliff and is from twenty to thirty yards
wide. On February 20 and 21, 1937 a large number of “Ele-
phant Trees,” Bursera microphylla, was observed spread over the
foothills on the east side of the Vallecito (Pifion) Mountains,
eastern San Diego County, for a distance of five or six miles
northwest of the gypsum mine near Split Mountain. It was es-
timated that there were at least two thousand trees in the region.
This species ranges from Arizona to Sonora and Lower Cali-
fornia but has been considered rare in the Colorado Desert.

Part one, volume two, of the “Flora of California” by Willis
Linn Jepson was issued on September 17, 1936. This fascicle,
consisting of pages 1 to 16 and 387 to 684, completes the volume

which includes the families from Capparidaceae to Cornaceae.

An historical sketch of descriptive floras for California from 1838
to 1880 introduces the volume; an index of families and genera
concludes it. Part two of volume two, consisting of pages 17 to
176, was issued February 15, 1936, and part three, pages 177 to
336, July 20, 19386. (Associated Students’ Store, University of
California, Berkeley, Unbound, $7.00; bound, $8.00.)

Dr. Yukio Yamada, a director of the Department of Botany,
Hokkaido Imperial University, Sapporo, Japan, has been in Berke-
ley since the latter part of January of this year. On February 26

1937] NOTES AND NEWS 71

he left for Los Angeles and from there will sail for Japan for the

_ opening of the college term in April. Dr. Yamada made a special

trip to this country, bringing with him collections of Sargassum
and Liagora for comparison with material in the algae collection
of the herbarium of the University of California at Berkeley.

Professor H. E. McMinn, of the Department of Botany, Mills
College, California, is taking advantage of his sabbatical leave
to further his knowledge of California shrubs by study of the
type specimens deposited in various herbaria in this country and
abroad. His itinerary from March 1 to August 1, 1937, includes:
Rancho Santa Ana Botanic Garden, Pomona College Herbarium,
and Blaksley Botanic Garden, Santa Barbara, in California; in
the east, Notre Dame University where Dr. E. L. Greene’s type
specimens are deposited, Gray Herbarium, and other leading bo-
tanical institutions; in Europe, Kew Gardens and the British
Museum, important botanical institutions on the continent, and
a month in Norway and Sweden. He hopes to study especially
the shrubs of northern Norway.

‘Dr. Mildred E. Mathias, who formerly worked at the Missouri
Botanical Garden, Carnegie Museum, and Pennsylvania State
College, recently moved to Berkeley and is continuing her studies
on the Umbelliferae at the Herbarium of the University of Cali-
fornia.

After nearly two and one-half years of botanical collecting in
South America, Mrs. Ynes Mexia disembarked at San Diego,
California, en route to San Francisco, on January 28, 1937.

She was in Ecuador from September, 1934, to September,
1935, collecting for the Bureau of Plant Introduction and Ex-
ploration of the United States Department of Agriculture certain
palms, cinchonas, and plants suitable for soil binders. Approxi-
mately five thousand herbarium specimens, as well as seeds and
bulbs, were secured from along the coastal plains, on the eastern
slopes of the Andes, and on the cold highlands of northern Ecua-
dor to the borders of Colombia.

From October, 1985, to January 1, 1986, Mrs. Mexia col-
lected for the University of California Botanical Garden Expedi-
tion to the Andes, which was engaged in obtaining native Nicoti-
ana species and plants of ornamental value. In addition to nearly
two thousand herbarium specimens, she secured seeds of practi-
cally all the Nicotiana species collected. During this period Mrs.
Mexia collected in Cerro de Pasco, Peru, then, returning to Lima,
went, by way of Lake Titicaca, to La Paz, Bolivia. After a col-
lecting trip into the hot provinces of Las Yungas, she left La Paz
and passed over the Bolivian highlands, through northern Argen-

"2 MADRONO [Vol. 4

tina and the warm plains of central Argentina to Tucuman, and
finally to Mendoza.

After terminating her connection with the California Botani-
cal Garden Expedition, Mrs. Mexia obtained approximately
twelve thousand herbarium specimens besides seeds and living
plants. These included, in addition to her Peruvian collections,
a representation of the antarctic flora of Tierra del Fuego, and of
the wet tropical vegetation of the province of Esmeraldas in
northwestern Ecuador.

PROCEEDINGS OF THE CALIFORNIA
] BOTANICAL SOCIETY

Wednesday, December 16, 1936. A meeting was held in
Room 2098, Life Sciences Building, University of California,
. Berkeley, at 8:00 p. m. The report of the nominating com-
mittee, read by Professor D. R. Hoagland, chairman, was as fol-
lows: president, Dr. F. W. Foxworthy; first vice-president, Dr.
Ira L. Wiggins; second vice-president, Dr. P. A. Munz; treasurer,
Dr. David D. Keck; secretary, Miss E. Crum. Dr. George J.
Peirce was nominated for president from the floor. Mr. Leo D.
Whitney gave an illustrated lecture on “Botanical Work in
Hawaii.”

Thursday, January 28, 1987. A meeting was held in Room
2098, Life Sciences Building, University of California, Berkeley,
at 8:00 p. m., Miss Alice Eastwood, first vice-president, presid-
ing. Dr. George J. Peirce having withdrawn his name, the officers
nominated at the previous meeting were unanimously elected.
Following the business meeting Dr. Peirce spoke on his recent
European tour.

Saturday, February 27, 19387. The annual dinner of the
California Botanical Society was held at the Hotel Durant,
Durant Avenue near Bowditch, Berkeley, at 6:30 p.m. Dr.
George J. Peirce, past president, introduced the new president
Dr. F. W. Foxworthy, chief forest research officer, retired, of
the Federated Malay States, now a resident of Berkeley. Dr.
Foxworthy then called upon Mrs. Ynes Mexia, Dr. Herbert L.
Mason, and Dr. David D.Keck to respond to toasts. A lecture
followed: “Domestication of plants under primitive cultures,” by
Dr. C. L. Alsberg of the Food Research Institute, Stanford Uni-
versity. Violin music offered under the direction of Mr. W. W.
Carruth, Mills College, was much appreciated. About sixty-five
members and guests attended the meeting.

Complete Your Files!

MADRONO

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“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges.

Volume I, 1916-1929... $5.00
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There is only a limited supply of Vol-
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NUMBER 3

MADRONO

- WEST AMERICAN JOURNAL OF
BOTANY

we
Contents othe

UPL ta mice ciate ng) (02

_ Published at Lime and Green Streets, Lancaster,
Pennsylvania

July, 1937

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asrams, Stanford University, California.

Dr. Lincotn Constance, University of California, Berkeley.

Dr. H. F. Coretann, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Dr. P. A. Munz, Pomono College, Claremont, California.

Business Manager—Dr. Davin D. Kreck
Lime and Green Sts., Lancaster, Pennsylvania
or
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Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

Established 1916. Published quarterly. Subscription Price $2.50 per year.
Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17,
complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num-
bers $0.75.

Papers up to 15 or 20 pages are acceptable. Longer contributions may
be accepted if the excess costs of printing and illustration are borne by the
contributor. Articles may be submitted to any member of the editorial board
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manuscripts may be included in the forthcoming issue provided that the
contributor pay the cost of the pages added to the issue to accommodate his
article. Reprints of any article are furnished at a cost of 4 pages, 50 copies
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Published at Lime and Green Streets, Lancaster,
Pennsylvania, for the

CALIFORNIA BOTANICAL SOCIETY

President: Dr. F. W. Foxworthy, Berkeley, California. First Vice-Presi-
dent: Dr. Ira L. Wiggins, Department of Botany, Stanford University. Sec-
ond Vice-President: Dr. Philip A. Munz, Department of Botany, Pomona
College, Claremont, California. Treasurer: Dr. David D. Keck, Carnegie In-
stitution of Washington, Stanford University. Secretary: Miss Ethel Crum,
4004 Life Sciences Building, University of California, Berkeley.

Annual membership dues of the California Botanical Society are $2.00,
$1.50 of which is for a year’s subscription to Madrofio. Dues should be
remitted to the Treasurer. General correspondence and applications for
membership should be addressed to the Secretary.

1937] BLAKE: TRACYINA "3

TRACYINA, A NEW GENUS OF ASTERACEAE
FROM NORTHERN CALIFORNIA

S. F. Biaxe

Dr. Herbert L. Mason of the University of California has
recently referred to me for examination specimens of a composite
collected in Humboldt County, California, in 1916 by Mr. Joseph
P. Tracy. The plant is superficially similar to Rigiopappus lepto-
cladus Gray, with which it occasionally grows, in stem and foliage,
and particularly in its habit of developing not far below the
terminal head a group of filiform 1-headed branches which surpass
the main stem and often branch similarly themselves. In its
technical characters, however, the plant shows a close relationship
to Pentachaeta, although it differs so strikingly from all members
of that genus in its slender-fusiform short-beaked achene, its
graduate pappus of numerous capillary bristles, and various minor
characters that it must be regarded as the representative of a new
genus. In Pentachaeta the involucre is 2-seriate and subequal, or
strongly graduated and about 5-seriate (in P. aurea), of mostly
oblong or lance-oblong, rarely lance-linear, thin and submem-
branous, greenish-centered, scarious-margined phyllaries, which
persist even after the fall of the achenes; the achenes are oblong
or obovoid, somewhat compressed, weakly 5-nerved or nerveless,
truncate, not at all narrowed or rounded and slightly contracted
at the apex; the pappus is variable, of 3-24 strictly 1-seriate
slender rigid hispidulous bristles, typically shortly paleaceous-
dilated at base, persistent or fragile and easily detergible, some-
times reduced to short bristles or cusps, or even wanting; and the
style branches have a short stigmatiferous portion and much
longer, linear, obtuse to acute, hispid or hispidulous appendages.
In the new genus the involucre is strongly graduate, about 4-
seriate, of narrowly linear readily deciduous phyllaries; the
achenes are slenderly fusiform, subterete, and weakly about 5-
nerved, drawn out above into a more or less distinct beak with
slightly dilated pappiferous apex; the distinctly graduated pappus
is composed of about 36-38 fragile but rather persistent, his-
pidulous, capillary bristles; and the stigmatiferous portion of the
style branches is about equal to the lance-subulate acuminate
hispid appendage. |

The “Herba Impia” habit of the new plant, so suggestive of
Rigiopappus, is rarely seen in Pentachaeta aurea but can be matched
in occasional examples of that plant, for example G. B. Grant 924
(Strawberry Valley, San Jacinto Mts., in U. S. Nat. Herb.). In
this collection the peduncles and tips of branches are glabrous at
apex, but in other specimens they are loosely pilose precisely as in
the new plant. The leaves of the new plant are a little broader

Maproxo, Vol. 4, pp. 73-104, July 1, 1937.

ny @ 1987

vey MADRONO [Vol. 4

and firmer and perhaps more stiffly ciliate than the average of
Pentachaeta aurea, but, considering the variation exhibited by this
species as a whole, I can find in herbarium specimens no constant
distinctive character until one comes to the heads. Somewhat

similar cases of “mimicry” are shown by other California Aster-,

aceae—particularly Crockeria chrysantha Greene and Lasthenia
glabrata Lindl., Monoptilon bellidiforme T. & G. and M. bellioides
(Gray) Hall (the two latter very distinct in their extreme forms,
but too intimately connected by apparently individual variation
for satisfactory generic separation )—but in these cases the ranges
of the members of each pair overlap. Pentachaeta aurea, how-
ever, is a plant of southern California, not known within about
three hundred miles of the area of the new plant, although other,
not “‘mimetic”’ species of Pentachaeta range north to Marin and
Mendocino counties. Mrs. Katherine Brandegee,’ discussing the

case of Crockeria and Lasthenia, suggested hybridity as the ex- »

planation of the origin of one of the forms in question, but in the
present case this explanation cannot be invoked. The one likely
parent is absent and no candidate for the place of the other is
evident. Pentachaeta aurea, Rigiopappus leptocladus, and the new
plant all occupy, apparently, much the same sort of habitat—dry
grassy hillsides—and their resemblances in habit and foliage can
probably be correlated with this fact, reinforced in the case of the
new plant by a probable genetic connection with the Pentachaeta
stock. The structural differences separating it from Pentachaeta
are, however, too significant to permit its inclusion in that genus.

Although the natural relationship of the new genus is un-
doubtedly with Pentachaeta, its much more copious and graduated
pappus would place it in the accepted arrangement of the sub-
tribe Solidagininae next to Aplopappus, from which its beaked
achene at once distinguishes it. The genus may appropriately be
named for its discoverer, Mr. Joseph P. Tracy of Eureka, Cali-
fornia, whose extensive collections in northwestern California,
made over a period of more than thirty years, have contributed
materially to our knowledge of the flora of that part of the state.
A nearly complete set of his collections, amounting to over twelve
thousand numbers, is deposited in the Herbarium of the Univer-
sity of California.

Tracyina Blake, gen. nov. Herba annua parva tenuissima
prope apicem pauciramosa praeter apices caulis ramorumque et
margines foliorum glaberrima, foliis alternis lineari-+lanceolatis
integerrimis hirsuto-ciliatis, capitulis mediocribus solitariis multi-
floris heterogamis flavis inconspicue radiatis, radiis erectis. In-
volucri subcylindrici (sice. turbinati) gradati ca. 4-seriati phyl-
laria linearia acuminata appressa plana 1-vittata submembranacea
viridescentia anguste scarioso-marginata. Receptaculum parvum
planum nudum scrobiculatum. Radii 1-seriati feminei fertiles

1 Zoe 5:95. 1901.

—

1937] BLAKE: TRACYINA 15

flavi dorso plusminusve rubicundi, tubo elongato, lamina parva
erecta elliptica bidenticulata. Flores disci flavi quoad numerum
radiis subaequales hermaphroditi fertiles, corollis tubulosis
anguste cylindrico-infundibuliformibus, tubo in faucem sensim
ampliato, dentibus 5 parvis ovatis erectis. Stamina 5, antheris
basi integris, apice appendice parva triangulari acuminata
munitis. Styli rami lineari-lanceolati, appendice lanceolato-
subulata acuminata hispida partem stigmatiferam aequante donati.
Achenia tenuiter fusiformia subteretia tenuiter ca. 5-nervia his-
pidula basi paullum angustata sursum longe angustata breviter
rostrata, apice ipso pappifero paullum dilatato. Pappi persistentis
sed fragilis gradati setae ca. 36-38 capillares hispidulae.—Species
typica T. rostrata, sp. nov.

Tracyina rostrata Blake, sp. nov. Annua erecta pedalis
saepius paene ad apicem simplex, prope apicem pauciramosa,
ramis filiformibus supra medium bracteatis prope apicem saepe
iterum ramosis; caulis glaber foliosus; folia alterna anguste lineari-
lanceolata acuminata sessilia integra erecta laete viridia 1-nervia
hirsuto-ciliata; capitula ca. 30—48-flora in apicibus caulis et
ramorum et ramulorum solitaria mediocria v. parva, primarium
pedunculatum cetera subsessilia; phyllaria anguste linearia
glabra; radii ca. 15-22, corollis ligulatis erectis disco paullo
longioribus; achenia radii sparsissime, ea disci dense hispidula;
pappus albidus achenio brevior.

Plant 15-30 cm. high; stems solitary, erect, whitish, usually
simple nearly to apex or occasionally with a few erect branches
from near the base, terete, not striate, 1 mm. thick or less, strictly
glabrous, terminated by a single head and usually bearing about
1.5—5 cm. below this head 2—4 filiform ascending branches 2.5—
10.5 cm. long, these arising close together (within 0.5-2 cm.),
naked below, rather densely leafy from about the middle to apex
with reduced, bract-like, subulate or lance-subulate leaves and
often branched in the same fashion as the main stem; leaves ap-
pressed or erect, narrowly lance-linear, the larger 1.2—2.4 cm.
long, 1-2 mm. wide, acuminate, acutely callous-apiculate, sessile
by a slightly narrowed base, entire, plane or somewhat involute
toward tip, light green, densely hirsute-ciliate, otherwise glabrous,
or obscurely hirsutulous on upper face, 1-nerved and with 1 pair of
weak lateral veins; terminal head usually on a more or less dis-
tinct, sparsely setaceous-bracteate peduncle 1.5—2.5 cm. long,
those of the branches usually sessile, the peduncle or branch some-
what thickened and striate below the head and there loosely
pilose; heads about 30—48-flowered, turbinate when pressed, sub-
cylindric when moistened, 9-10 mm. high (in fruit), the larger
1.5 cm. thick (in fruit, as pressed) ; involucre 6—7 mm. high, the
phyllaries narrowly linear (0.8-0.5 mm. wide), passing into the
bracts of the branch, readily deciduous; rays about 15—22, the tube
about 3 mm. long, glabrous or slightly hispidulous above, the

"6 MADRONO [Vol. 4

Fig. 1. Tracyina rostrata Blake. a, upper part of plant, x1; b, disk
achene, X 8; c, disk flower, <8; d, style branches, xX 28; e, two stamens, X32;
f, ray flower, x 8; g, head, x3; h, ray achene, <8, part of the pappus omitted. |
All drawn from the type.

lamina narrowly elliptic, about 1.5 mm. long, 0.4 mm. wide,
bidenticulate, glabrous, in the dried state pale yellow, reddish on
the back; disk flowers about 15-26, pale yellow, sparsely hir-
sutulous on throat and teeth, slenderly cylindric-funnelform,
4 mm. long (tube 1.4 mm., throat 2.3 mm., teeth ovate, 0.38 mm.
long); mature achenes slenderly fusiform, shortly rostrate, not
filled by the seed, 5.5 mm. long, 0.3-0.4 mm. thick, those of the

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 17

ray very sparsely, those of the disk densely hirsutulous with
bidenticulate hairs; pappus of about 36-38 graduated persistent
hispidulous bristles, the inner 3.8 mm., the outer about 1 mm. long.

Cauirornia. Dry grassy hills, Alder Point, alt. 245 m., 23
May 1903, Tracy 1892 (Herb. Univ. Calif.); on warm grassy
slopes, Alder Point, on Eel River, southeastern Humboldt Co.,
alt. 8300 m. (1000 ft.), 20 June 1916, J. P. Tracy 4735 (type no.
549767, Herb. Univ. Calif.; duplicate, U. S. Nat. Herb.) ; grassy
slopes, Dobbyn Creek, Humboldt Co., alt. 150 m., 15 June 1930,
Tracy 8762 (U.S. Nat. Herb.)

According to information furnished by Mr. Tracy, the plant is
frequent but inconspicuous in short grass on warm grassy slopes
at elevations of 300 to 1000 feet from about two miles south of
Alder Point northeast about five miles to Dobbyn Creek. It is
very similar in habit to Rigiopappus leptocladus, but taller, and
generally grows in better soil, although occasionally the two are
found together. The fruiting heads resemble those of Agoseris
heterophylla, which grows abundantly in the same places.

Bureau of Plant Industry, Washington, D. C.
March, 1937.

VEGETATIONAL SURVEY OF THE LOWER
RIO GRANDE VALLEY, TEXAS

Exvzapa U. CLover

(Concluded from page 66, issue of April 7, 1937)
III. Coastat Criimax ASSOCIATIONS

This area includes the sandy beach, sand dunes (medaios),
salt flats (badilla salitrosa) and the salt grass region known as the
sacahuistal. There are certain differences in the types of vege-
tation at different points along the coast. At Red-Fish Bay
(Willacy County) where the Sand Belt dips down to the coast,
the zacatal vegetation persists to the water’s edge. Some Spar-
tina Spartinae may be found, but in general the typical coast flora
is lacking. Just south of the Bay, long estuaries reach inland
for some distance and the Raymondville and Port Isabel Clays
support the flora common to salt marshes.

Boca Chica offers an excellent opportunity for a study of
coast phytogeography. The beach is sandy with shifting dunes.
Back of these low dunes there are others sometimes twenty to
twenty-five feet high which are stable and covered with vegeta-
tion. Barren salt flats and salt marshes reach out into the grass-
land along the coast, and small islands lie between Brazos Santi-
ago Island and the mainland.

Beacu VecetTation. There are few plants along the beach
which are not found on the dunes. In fact the dunes usually are
reached by high tides. Cakile maritima var. aequalis, Portulaca

78 MADRONO | [Vol. 4.

pilosa, and Oenothera Drummondii are found on sandy flats between
dunes. |
Surrtinc Dunes. Uniola paniculata, Fimbristylis castanea, Ipo-
moea Pes-Caprae, and Sesuvium verrucosum serve as sand binders on
the shifting dunes.
PerMANENT Dunes. These dunes are covered by a prairie

phase of the coastal vegetation. A list of plants found on per--

manent and shifting dunes at Boca Chica is listed below: Di-
stichlis spicata, Spartina Spartinae, Eleocharis albida, Sisyrinchium
longipedunculatum, Batis maritima, Suaeda conferta, D. multiflora,
Atriplex matamorensis, Cladothriz lanuginosa, Philoxerus vermicularis,
Portulaca oleracea, Sesuvium verrucosum, Cakile americana, Hoff-
manseggia densiflora, Indigofera leptosepala, Mimosa strigillosa, Petal-
ostemon emarginatus, Strombocarpa cinerescens, Samodia ebracteata,
Abutilon incanum, Oenothera Drummondii, Limonium Nashi, Sabbatia
carnosa, Podostemma sp., Ipomoea Pes-Caprae, Ipomoea stolonifera,
Opuntia Allairei, Agalinis heterophylla, Aplopappus phyllocephalus,
Borrichia frutescens, Gaillardia pulchella, Erigeron repens.

Tue Sart-ryat. Salt flats which are covered with water by
gulf storms are almost barren. Most vegetation occurs near the
margins with a few straggling plants, mostly of Salicornia, invad-
ing the salt-incrusted flat. Paspalum distichum, Monanthochloé
littoralis, and Chamaecrista littoralis venture beyond the margins
and an occasional sand dune out in the barren flat has the usual
dune vegetation.

Low Istanps. Hurricanes are of infrequent occurrence, some-
times doing little or no damage to the south Texas coast for
periods of ten or fifteen years. The flora farther inland becomes
fairly well established. Minor storms keep the vegetation of the
salt flats and marshes and some of the low islands in stages of
both progressive and retrogressive succession (8, 15). Clark
Island which lies one mile west of the northern end of Brazos
Island is an exception. It is somewhat higher, at least parts of
it, than the others, and is also protected by Brazos Island, thus

escaping the violent action of the waves. This island is roughly |

elliptical in shape, measuring about one mile in length and one-
half mile across. ‘The north end is several feet above water-level,

and is covered with mesquite trees which are ten to twelve feet ©

high. This is a nesting place for herons, and the trees are
weighted down with bulky masses of sticks which constitute the
nests. A single specimen of Yucca Treculeana measuring ten feet

grew on the island, but was blown down in the hurricane of 19338.

About one-third of the island is covered by a coastal prairie and
upland vegetation, and the southeast portion is low and covered
with water in places. The southeast margin is bordered by a
low growth of Avicennia nitida (black mangrove). Salicornia,
Atriplex and Suaeda inhabit the wet area. It is interesting to find
a coastal flora such as that of Avicennia nitida with gradations to

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 79

Figure 2

Prate XVI. Fig. 1. Lophophora Williamsii growing on the Bordas
Scarp eight miles north of Rio Grande City, Starr County. This is the cactus
used for religious purposes by the Indians. Fig. 2. Ferocactus hamata-
canthus. 'This cactus is fairly abundant along the Rio Grande near Zapata.

80 - MADRONO [Vol. 4

the semi-desert vegetation. A complete list of the island flora
is given below: Andropogon littoralis, Aristida purpurea, Cenchrus
pauciflorus, Chloris Petraea, Eragrostis secundiflora, E. sessilispica,
Monanthochloé littoralis, Polypogon monspeliensis, Spartina Spartinae,
Sporobolus argutus, Vaseyochloa multinervosa, Cyperus uniflorus, Ele-
ocharis albida, Commelina crispa, Yucca Treculeana, Batis maritima,
Cladothrix lanuginosa, Atriplex matamorensis, Suaeda linearis, Sali-
cornia ambigua, S. herbacea, Lepidium virginicum, Baptisia leucophaea,
Dolicholus americanus, Hamosa Nuttalliana, Indigofera leptocephala,
Prosopis juliflora var. glandulosa, Sophora tomentosa, Strombocarpa
cinerescens, Kallstroemia parviflora, Polygala alba, Croton ciliato-
glandulosus, Galpinsia tubicula, Sabbatia carnosa, Opuntia Alleirei,
Avicennia nitida, Verbena Hallii, Physalis mollis, P. viscosa, Calceo-
laria verticillata, Solanum triquetrum, Stemodia lanata, Plantago rho-
dosperma, Aphanostephus skirrobasis, Baccharis texana, Coreopsis
cardaminifolia, Gaillardia pulchella, Othake robustum.

Tue Sacanuistat. This area extends several miles inland.
The dominant is Spartina Spartinae, in the marshes associated
with Suaeda, Salicornia, and sedges, and farther inland by Eupa-
torium odoratum,; Aplopappus phyllocephalus, Franseria confertifolia,
Pluchea camphorata, Senecio glabellus, Setaria macrostachya, and
Sporobolus argutus. Mesquite trees invade this coastal grass area,
with Opuntia and some Yucca. Finally shrubs, especially Lycium
carolinianum, Zizyphus obtusifolia, Celtis pallida, and Forestiera an-
gustifolia help form the transition zone between the sacahuistal
and the mesquital-chaparral.

Yucca Treculeana occurs as “forests” near Boca Chica and
Port Isabel. The hurricane of 1933 completely wiped out this
species at Boca Chica.

BoscasE pE Patma. Probably the most picturesque and cer-
tainly one of the most interesting associations in the Lower Rio
Grande Valley is the native palm grove ten miles below Browns-
ville (pl. XVII). Rio de las Palmas was the first name for the
river later known as Rio Bravo del Norte (17), and now called
EI Rio Grande. This first name is found on an old Spanish map
(Cantino, 1502) of North America, perhaps the first one made
(31). No doubt the name was suggested by the native palms
which grew on its banks. This palm, Sabal texana, la palma de
micharos, is one of the four arborescent palms occurring in the
continental United States outside of Florida. The others are the
delta palm, Sabal Deeringiana, of the Mississippi delta, the cab-
bage tree, Sabal Palmetto, of the southeastern Atlantic region, and
the fan-leaf palm, Neowashingtonia robusta, of the California
deserts.

Arthur V. Schott, a geologist and surveyor in the Mexican-
United States Boundary Survey, collected plants in the Lower
Rio Grande Valley, and probably made the first report of any
importance regarding these palms. The following is a quotation

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 81

Figure 2

Puate XVII. Fig. 1. Sabal texana. This palm extends over approxi-
mately one hundred acres along the Rio Grande below Brownsville. Fig. 2.

Cordia Boissieri. When land is cleared native trees are sometimes allowed to
remain.

82 MADRONO [Vol. 4

from his report (29): “It is also in the lower portion of this belt
[coastal plain of Texas] where the palm tribe is represented by
Chamaerops Palmetto that the palmetto attains a growth as gor-
geous even as that on the lower Mississippi; it extends on the
Rio Bravo up to about eighty miles from the Gulf.”

Sabal texana in the United States is confined to a limited area
in Cameron County near the Rio Grande extending from El
Salado ranch ten miles below Brownsville up the river three or
four miles, forming a heavy growth. This subtropical forest with
its tangle of vines, shrubs, and fallen trees is almost impenetra-
ble. The palms grow to the height of fifty feet on the Rabb
ranch where the finest growth occurs. This palm has often been
confused with Sabal mexicana of southern Mexico. Arundo Donaz
is twenty feet tall here and adds to the tropical appearance of
the locality, as does Malvaviscus Drummondii. It is difficult to tell
what plants rank second or third in dominance, but prominent
ones are: Ulmus crassifolia, Pithecolobium brevifolium, Celtis missis-
sippiensis, Celtis pallida, Fraxinus Berlandieri, Leucaena pulverulenta,

Tillandsia usneoides, and Arundo Donax. Below is a list of plants

collected from the river’s edge into the densest palm growth:
Sabal texana, Arundo Donaa, Echinochloa colonum, E. Crusgalli, Era-
grostis hypnoides, Oplismenus setarius, Panicum fasciculatum, P.
purpurascens, Paspalum conjugatum, Sporobolus Buckleyi, Carex Brit-
toniana, Cyperus ochraceus, Tillandsia recurvata, T. usneoides, Com-
melina longicaulis, Smilax renifolia, Celtis mississippiensis, C. pallida,
Ulmus crassifolia, Persicaria longistyla, P. mexicana, Rumezx Ber-
landiert, Acanthochiton Wrightw, Amaranthus hybridus, A. spinosus,
Rivina humilis, Clematis Drummondii, Roripa Nasturtium, Rubus trivi-
alis, Erythrina herbacea, Mimosa Berlandieri, Leucaena pulverulenta,
Pithecolobium brevifolium, Ionoxalis sp., Xanthoxylum insulare, X.
Pterota, Malphigia glabra, Chamaesyce hypericifolia, Poinsettia heter-
ohpylla, Tragia nepetaefolia, Cardiospermum corindum, C. Halica-
cabum, Zizyphus obtusifolia, Ampelopsis arborea, Abutilon Jacquinii,

Malachra urens, Malvastrum spicatum, Malvaviscus Drummondii, Wis-,
sadula amplissima, Gaura parviflora, Passiflora lutea, Plumbago scan- |

dens, Bumelia angustifolia, Diospyros texana, Forestiera angustifolia,
Fraxinus Berlandieri, Metastelma barbigerum, Cuscuta arvensis, Pha-
celia patuliflora, Lantana horrida, Lippia alba, Verbena (hybrid),
Salvia Greggu, Solanum nigrum, Melothria pendula, Ammannia cocei-
nea, Centaurea americana, Eclipta alba, Erigeron tenuis, Eupatorium
odoratum, Parthenium Hysterophorus, Senecio glabellus, Verbesina
virginica.
SUCCESSION IN CULTIVATED AREAS

Much of the land below Pefiitas has been cleared for several
miles back from the river and is under cultivation. Irrigation
projects have been responsible for much of this clearing. Some
of this land has been abandoned. Primary succession in an un-

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 83

irrigated area near Pefiitas is characterized largely by the follow-
ing species: Panicum firmulum, P. Halli, Paspalum Langei, Chamae-
syce cordifolia, Croton leucophyllus, Solanum elaeagnifolium, Verbena
Halei, Ambrosia elatior, Parthenium Hysterophorus, and Verbesina
encelioides. This is usually followed by Celtis pallida, Zizyphus
obtusifolia, Prosopis juliflora var. glandulosa, and Opuntia Lind-
_heimeri. Near the river Clematis Drummondu, Amaranthus hybridus,
Cuscuta arvensis, Cardiospermum Halicacabum, Solanum triquetrum,
and Parthenium Hysterophorus are some of the first plants to in-
vade an abandoned clearing. In the irrigated section Cenchrus
pauciflorus, Cynodon Dactylon, Holcus halepensis, Setaria macro-
stachya, Amaranthus, and Parthenium Hysterophorus are most
important during primary succession.

In many places native trees have been permitted to stand in
clearings; and when the land is abandoned, seeds from these
trees start new colonies of the former vegetation. Some of these
favored species are: Cordia Boissieri, Siderocarpos flexicaulis, Dio-
spyros texana, and Prosopis juliflora var. glandulosa.

RELATIONS oF Epapuic Factors AND GEOLOGICAL FORMATIONS
To Pirant DistTRIBUTION

A comparison of the geological map (text fig. 1) and the
vegetation map (text fig. 3) shows that in many instances there
is a close correlation between the geologic formations and plant
distribution in the Lower Rio Grande Valley.

The zacatal or prairie grass region conforms almost exactly to
the wind-blown sands and beach deposits.

In general, the mesquital-zacatal in Zapata County (which is
included in the Aguilares Plain) conforms to the Fayette Sand-
stone formation. In places the surface materials are clay and
sand with gravel from the Reynosa formation and consequently
areas of chaparral appear in places throughout the mesquital-
zacatal.

The Reynosa formation covering much of Starr County and a
strip along the west side of Hidalgo County is largely mesquital
divided into the same three phases which continue toward the
coast. However, there is a difference in the appearance of the
brush. The growth here tends to be smaller and sparser with
fewer mesquites present.

The Bordas Scarp begins at Rio Grande City and continues
through Starr County to Cuevitas. It extends northward into
Jim Hogg County and cuts the northeast corner of Zapata
County. The vegetation along this escarpment consists of a
sparse growth of chaparral with more species of cacti than are
found on either side of this elevation.

Terrace deposits in general produce a growth of large mes-
quites. This is well shown at San Ignacio and Sam Fordyce.

84 MADRONO [Vol. 4

There is little difference in the vegetation on Beaumont clay
and the so-called ‘‘Fluviatile’’ deposits except toward the coast.
The chaparral here shows a more luxuriant growth than that of
the Reynosa.

There is no doubt that moisture is an important limiting fac-
tor. The vegetation along the river in the lower part of this
region is composed of species which require more moisture than

those in the upper valley. Some species which grow well near_

the coast are hardly recognizable for their diminutive size in dry
gravel areas.

Salt depressions have characteristic beach and coastal prairie
vegetation at their margins, although a few yards distant the in-
land type of plant growth is dominant. Poorly drained soil has

become so impregnated with alkali salts as to become practically ©

useless for farming. This condition naturally limits all vegeta-
tion to alkali resistant plants such as the salt grasses, and Suaeda,
Varilla texana, Salicornia, and Lycium carolinianum.

Hechtia texensis seems to prefer soil of the type found in the
Cockfield formation. This formation was deposited in salt and
brackish marshes and must still be at least slightly saline since
Varilla texana (a halophyte) is always associated with it. Dry
gravel hills are covered with Acacia amentacea often accompanied
by Acacia Berlandieri, Leucophyllum frutescens, Mortonia Greggiu
and Cordia Boissieri, species which ordinarily prefer limestone
soil. More species of cacti are found in this Lissie Gravel than
in any other formation. This may be either because they are
better adapted to this soil, or because there is a sparse growth of
brush, thus permitting them to have better light conditions.

The most luxuriant vegetation is in the alluvial soil along the
Rio Grande below Pefiitas. As moisture increases toward the
coast the vegetation increases in size and number of species.
Plants with tropical affinities are more abundant in the region of
Brownsville probably because of better moisture conditions and
slightly higher temperature. The soil is more favorable for
growth, allowing chance seeds to germinate.

EFFECT oF CHANGES IN PHyYSIOGRAPHY

The changing course of the Rio Grande causes changes in
vegetation. Alluvial soil is deposited in places and carried away
in others. Floods fill up resacas, killing some plants and per-
mitting the growth of others. Gulf storms destroy vegetation by
wind-action and by blowing salt water inland. The filling of
estuaries causes very unstable conditions for plant development.
Arroyos dig deeper and wider with each succeeding storm, in-
fluencing the vegetation along the banks. Water-holes are
formed in the beds of arroyos allowing such trees as Acacia Far-
nesiana and Parkinsonia aculeata to become established on the
banks.

1937 | CLOVER: VEGETATION OF RIO GRANDE VALLEY 85

ACKNOWLEDGMENTS

I wish to express appreciation to Dr. E. J. Alexander, Assis-
tant Curator at the New York Botanical Gardens, for determin-
ing and checking much of the material; to Dr. S. F. Blake,
National Herbarium, Smithsonian Institution, who determined
most of the Compositae; to Dr. Hugh O’Neill, Catholic Univer-
sity, Washington, D. C., who determined the Cyperaceae; and to
Dr. William C. Steere who determined the Hepaticae and Musci.
The late Dr. A. S. Hitchcock, Department of Agriculture, Bureau
of Plant Industry, determined the Gramineae. I am much in-
debted to Professor H. H. Bartlett and to Dr. J. H. Ehlers for
their helpful advice in the study of the problem and the prepara-
tion of this paper, and to the Department of Botany and the
Botanical Gardens of the University of Michigan for financial
assistance in the field survey.

Species COLLECTED IN THE LOWER Rio GRANDE VALLEY?

RICCIACEAE

OxYMITRA ANDROGYNA Howe. Dry, red, sandy soil, Rio Grande City.
Riccra sp. On sandy-loam soil, mesquite woods, Rio Hondo.

LEJEUNEACEAE
FRULLANIA squarRosa (R., Bl. & N.) Dumort. On living branches of
Castela Nicholsontit, Raymondville.
FRULLANIA EBORACENSIS Gottsche. Mesquite woods, Rio Hondo.

PorrraceaE
PTeRYGONEURUM cCaAviroLtiIum (Ehrh.) Jur. On dry, sandy loam, Rio
Grande City.
TRICHOSTOMUM BRACHYDONTIUM Bruch. On dry, sandy-clay soil, Zapata.
Wetsta AnpDrewsit Bartr. On limestone rock in full sunlight, La Joya.

FUNARIACEAE

PyRAMIDULA TETRAGONA (Brid.) Brid. On sandy soil in mesquite woods,
Raymondville.
PHYSCOMITRIUM TURBINATUM (Michx.) Brid. On damp alluvial soil, road-

.Side, Barreda.

PHYSCOMITRIUM IMMERSUM Sulliv. In deep cracks made by mud drying
after flood, Pefitas.

Bry ACEAE

BryuM ARGENTEUM Hedw. var. LANosum Bryol. eur. Dry sandy soil, in
low brush, Raymondville.

ERPODIACEAE

ERPODIUM DOMINGENSE (Brid.) C. Miill. In shade, on decaying wood, Rio
Hondo.

THUIDIACEAE

HapLociaDIUM MICROPHYLLUM (Hedw.) Broth. On decaying log in moist
woods, Palm Grove, Brownsville.

3 Numbers in parentheses refer to collections by the author.

86 MADRONO [Vol. 4

PLAGIOTHECIACEAE

STEREOPHYLLUM Wricutt (Sulliv.) Ren. & Card. On decaying: log in
moist woods, Alamo.

MaARSILEACEAE

MarsitEA mMacropopa Engelm. Progresso (18).
Marsitea vestiTa Hook. & Grev. Mission (115).

FILIcINEAE
NorHOLAENA DEALBATA (Pursh) Kunze. Roma (791).

PINACEAE
TaxoDIUM MUCRONATUM Tenore. Havana (1123).

GNETACEAE

EPHEDRA ANTISIPHILITICA Meyer. Edinburg (1593).
EPHEDRA PEDUNCULATA Engelm. Barrocetes Ranch, San Ignacio (1584).

POTAMOGETONACEAE
ZOSTERELLA DUBIA (Jacq.) Small. La Joya Lake (640).

TYPHACEAE
Typua tatirotia L. Chihuahua (214).

N AIADACEAE
NAIAS GUADALUPENSIS (Spreng.) Morong. Alamo (894).

ALISMACEAE

Ecuinoporvus corpirotius (L.) Griseb. Alamo (639).
SAGITTARIA VARIABILIS Engelm. Alamo (265).

GRAMINEAE

ANDROPOGON BARBINODIS Lag. Laredo (1278).

ANDROPOGON GLOMERATUS (Walt.) B.S. P. Weslaco (462).
ANDROPOGON LITTORALIS Nash. Padre Island (1633).
ANDROPOGON SACCHAROIDES Swartz. Edinburg (440).

ARISTIDA PURPUREA Nutt. Pefiitas (575).

Arunpbo Donax L. Palm Grove, Brownsville (959).
BoutTeLovua BARBATA Lag. Alamo (8).

BovutEetova Hirsuta Lag. San Ignacio (1586).

BovureLova TRiIFIDA Thurb. Salinefio (1406).

BRACHIARIA CILIATISSIMA (Buckl.) Chase. Monte Cristo (927).
Cencurus Incertus M. A. Curtis. San Juan (305).

CENCHRUS PAUCIFLORUS Benth. La Joya (586).

CHLORIS ANDROPOGONOIWES Fourn. Mission (162).

CuHLoris cucuLtata Bisch. Tabasco (91).

Cutoris Gayana Kunth. Donna (625).

Cuuoris Petrara Swartz. Clark Island (704A).

Cynopon Dacryton (L.) Pers. Alamo (433).
Dacrytocrentum Arcyptium (L.) Richt. San Juan (107).
DiIGITARIA SANGUINALIS (L.) Scop. Harlingen (1211).
Disticuuis spicata (L.) Greene. Boca Chica (383).
EcHINOCHLOA coLtonum (L.) Link. Chihuahua (985).
EcuinocHioa Cruscatyi (L.) Beauv. Palm Grove, Brownsville (1518).
EcHINOCHLOA CRUSGALLI ZELAYENSIS (H. B. K.) Hitche. Alamo (289).
ELEUSINE InpIcA (L.) Gaertn. Alamo (1350).

Eracrostis BARRELIERI Daveau. San Juan (451).

1937 |

CLOVER: VEGETATION OF RIO GRANDE VALLEY

Eracrostis Beyricui J. G. Smith. Cuevitas (1095).
ERAGROSTIS CURTIPEDICELLATA Buckl. La Joya Lake (43).

ERAGROSTIS HYPNOIDES (Lam.) B.S. P. Palm Grove, Brownsville on

Eracrostis tuGENS Nees. Kingsville (837).
ERAGROSTIS SECUNDIFLORA Presl. Boca Chica (720).
ERAGROSTIS SESSILISPICA Buckl. Clark Island (740).
ERAGROSTIS REPTANS (Michx.) Nees. Alamo (482).
EraGrosTis spicata Vasey. Alamo (297).

EriocHLoa PpunctTATA (L.) Desv. Brownsville (1650).
HeETEROPOGON conTortTus (L.) Beauv. Linn (847).
Hotcus HALerensis L. Alamo (149).
Lreptotoma cocnatum (Schult.) Chase.
Leprocutoa pusia (H. B. K.) Nees. Alamo (1481).
LEPTOCHLOA FILIFORMIs (Lam.) Beauv. San Juan (559).
LreprocHitoa NEaLLeyi Vasey. Rio Hondo (1580).

LEPTOCHLOA UNINERVIA (Presl) Hitche. & Chase. Alamo (918).
MoNANTHOCHLOE LITTORALIS Engelm. Clark Island (759).
OPLISMENUS sETARIUS (Lam.) Roem. & Schult.

Boca Chica (727).

(1508).

PANICUM FASCICULATUM Swartz. Palm Grove (956).

PaNnicuM FIRMULUM Hitche. & Chase. La Joya (580).

Panicum Harum Vasey. La Joya (557).

Panicum uians Ell. Encino (836).

Panicum nopatum Hitche. & Chase. Encino (834).

PANICUM PURPURASCENS Raddi. Palm Grove, Brownsville (1511).
PANICUM SPHAEROCARPON Ell. Encino (858).

Panicum TExANUM Buckl. Alamo (1347).

PaproPHoruM sicotor Fourn. Arroyo el Tigre (697).
PAPpPOPHORUM MUCRONULATUM Nees. San Manuel Ranch (851).

PasPaALUM
PasPpaALuUM
PasPALUM
PasPaALUM
PasPpALUM
PasPALUM
PasPaALUM
PHALARIS

PHRAGMITES COMMUNIs Trin.
POLYPOGON MONSPELIENSIS (L.) Desf.

consucAtum Berg. Palm Grove, Brownsville eet ve
DILATATUM Poir. Alamo (1349).

pisticHum L. Boca Chica (365).

Lancet (Fourn.) Nash. Tabasco (781).

Livipum Trin. Alamo (1115).

STRAMINEUM Nash. Encino (860).

Urvitter Steud. Donna (1014).

CAROLINIANA Walt. Encino (832).

Mercedes (630).

Clark Island (758).

SETARIA MAcROsTACHYA H. B. K. Rio Hondo (613).

.SETARIA SCHEELEI (Steud.) Hitchc.
SeTar1a setosa (Swartz) Beauv.
SETARIA GENICULATA (Lam.) Beauv.

SPARTINA

Alamo (1453).
Alamo (1453A).
Encino (839).

SparTInaE (Trin.) Merr. Boca Chica (729).

Sporopotus arcutus (Nees) Kunth. Clark Island (1256).
Sroropotus BucKkiEy1 Vasey. Palm Grove, Brownsville (1560).
SPoroBoLus cRYPTANDRUS (Torr.) Gray. Harlingen (1213).
SrorospoLus Wricutm (Munro) Scribn. Boca Chica (402).
TRICHACHNE CALIFORNICA (Benth.) Chase. Mission (1993).
TRICHACHNE SACCHARATA (Buckl.) Nash. Mission (174).
TRICHLORIS MENDOCINA (Phil.) Kurtz. Arroyo Velefio (1290).
TRICHLORIS PLURIFLORA Fourn. Weslaco (628).

TRICHOLAENA ROSEA Nees. Donna (466).

TRICHONEURA ELEGANS Swallen. San Ignacio (1582).

TRIODIA ALBESCENS Vasey. Harlingen (1025).

Triopia muTicA (Torr.) Scribn. Salinefio (1405).

Triop1a prtosa (Buckl.) Merr. Arroyo el Tigre.

Uniota panicuLtata L. Boca Chica (386).
VASEYOCHLOA MULTINERVOSA (Vasey) Hitchce.

Clark Island (1719).

87

Palm Grove, Brownsville

88

MADRONO [Vol. 4

CYPERACEAE

Carex Brirroniana Bailey. Progresso (18).

CyYPERUS
CYPERUS
CYPERUS
CYPERUS
CYPERUS
CYPERUS
CYPERUS
CYPERUS
CYPERUS
CyYPERUS
CyYPERuUS
CYPERUS

ELeocHARIS ALBIDA Torr.
ExLeocuaris pALustTRis (L.) Roem. & Schult.

acuminatus Torr. & Hook. Chihuahua (598).
articuLtatus L. Alamo (1119).

cyLtinpricus (Ell.) Britton. Ojo de Agua (995B).
ELEGANS L. Rio Hondo (1020).

ERYTHRORRHIZOS Muhl. Palm Grove, Brownsville (1529).
FERAX L. C. Rich. Progresso (15).

cLososus Aubl. Monte Cristo (929).

ocHRAcEUs Vahl. Palm Grove, Brownsville (1981).
oxycariowes Britton. Alamo (995).

stricosus L. Alamo (898).

uniFLorus Torr. & Hook. Alamo (150).

virENS Michx. Rio Hondo (1030).

Clark Island (747B).

Rio Hondo (1012).

FIMBRISTYLIS CASTANEA (Michx.) Vahl. Boca Chica (771).

ScIRPUS AMERICANUS Pers.
SciIrRPUS CALIFORNICUS Britton.

Boca Chica -(701).
Brownsville (973).

Scmpus Hatim Gray. Alamo (887).

Scirpus vauipus Vahl.

SaBaL TEXANA (O. F. Cook) Small.

La Feria (633).

PALMAE
Brownsville (1504).

LEMNACEAE

Lemna minor L. Rio Hondo.

WoOLFFIA COLUMBIANA Karsten. Rio Hondo.
|
BROMELIACEAE q

HecurTia TEXENSIS Wats. Roma (523).

TILLANDsIA Barttey1 Rose.

Alamo (1422).

TILLANDsIA RECURVATA L. La Joya (69).

TILLANDSIA usNEoIES L. Tabasco (226).

CoMMELINA cRISPA Wooton.
CoMMELINA LONGICAULIS Jacq. Alamo (1422).
SITREAVSIA PULCHERRIMA Bartlett.

HETERANTHERA LImMosA (Swartz) Willd. Rio Hondo (1038). |
EIcHoRNIA CRASSIPES Mart. |

JUNCUS ARISTULATUS Michx.

ALLIUM ARENICOLA Small.
Autium scarposum Benth. Boca Chica (1709). I
SMILAX RENIFOLIA Small.
Yucca rupicouta Scheele.

YUCCA TENUISTYLA Trel.

Yucca TRECULEANA Carr.
Yucca sp. Pefiitas (135).

AGAVE MELLIFLUA Trel.
AGAVE LEcHUGUILLA Torr.
AGAVE sp. Trel.

COMMELINACEAE
Roma (161).

Port Isabel (974).

PONTEDERIACEAE

Brownsville (361).

J UNCACEAE |
Raymondville (833). |

LILIACEAE
Arroyo el Tigre (1797).

Tabasco (227).

San Manuel Ranch (820).
San Ignacio (505).

Boca Chica (989).

AMARYLLIDACEAE

El Conchito Ranch, Zapata (1295).
Roma (477).
Rio Grande City (284).

v

a

e

1937 | CLOVER: VEGETATION OF RIO GRANDE VALLEY

Aramosco TExANA (Herb.) Greene. Alamo (1421).
Coorer1a Drummonpit Herb. San Juan (4).
Mawnrrepa Macutosa (Hook.) Rose. La Joya (62).
MANFREDA VARIEGATA (Jacobi) Rose. Alamo (12).
RuNYONIA LONGIFLORA Rose. Rio Grande City (483).

IRIDACEAE

SisyRINCHIUM AMOENUM Bicknell. Padre Island (1640).
SisyRINCHIUM FURCATUM Bicknell. San Manuel Ranch (808).

SISYRINCHIUM LONGIPEDUNCULATUM Bicknell. Boca Chica (707).

ZINGIBERACEAE
Crorria parvirotia (Torr.) Small. Edinburg (1587).

SALICACEAE
Satix toncirotIA Muhl. Alamo (261).

BATIDACEAE

a (Bans mariTiMA L. Clark Island (764).

FAGACEAE
QUERCUS VIRGINIANA Mill. N. Willacy County (638).

ULMACEAE

CELTIS MISSISSIPPIENSIS Bosc. Palm Grove (457).
Certis pattipa Torr. La Joya (75).
Uxmus crassiroLria Nutt. Tabasco (224).

URTICACEAE

PaRIETARIA OBTUSA Rydb. Alamo (1780).
Urtica CHAMAEDRYOIDES Pursh. Alamo (1863).

LoraNTHACEAE
PHORADENDRON FLAVESCENS Nutt. Rio Grande City (525).

ARISTOLOCHIACEAE

ARISTOLOCHIA LONGIFLORA Engelm. & Gray. Arroyo el Tigre (751).

POLYGONACEAE

ErRI0GONUM MULTIFLORUM Benth. Boca Chica (1913).
PERSICARIA LONGISTYLA Small. Palm Grove, Brownsville (1514).
PERSICARIA MEXICANA Small. Palm Grove, Brownsville (1455).
PERSICARIA PENNSYLVANICA (L.) Small. Alamo (903).

Rumex Berianpiert Meisn. Palm Grove, Brownsville (1530).

_-Rumex crispus L. Weslaco (608).

a =

|

6 aed

RumMex MeExicanus Meisn. Tabasco (166).
RumeEx sprrauis Small. Havana (1127). ~

CHENOPODIACEAE

ATRIPLEX ACANTHOCARPA (Torr.) S. Wats. Roma (1090).
ATRIPLEX ARENARIA Nutt. Port Isabel (1263).

89

ATRIPLEX CANESCENS (Pursh) Nutt. El Jaral Ranch, Zapata Co. (1581).

ATRIPLEX MATAMORENSIS Nels. Boca Chica (374).
CHENOPopDIUM ALBUM L. Boca Chica (1710).
CHENOPOoDIUM virIDE L. San Benito (1929).

, Donp1a conrerta Small. Boca Chica (354)

| Donpia trnearis (EIl.) Millsp. Port Isabel (1262).
Donpia MuLTIFLoRA (Torr.) Heller. Boca Chica (377).
Donpia sp. Clark Island (329). Possibly a new species.
SALIcoRNIA AMBIGUA Michx. Clark Island (767).

ao

90

MADRONO [Vol. 4

Saricornia BiceLtovit Torr. Boca Chica (359). |
SALICORNIA HERBACEA L. Clark Island (766).

AMARANTACEAE

AcaNnTHOCHITON Wricutu Torr. Palm Grove, Brownsville (1535).
ACHYRANTHES REPENS L. Boca Chica (395).

AMARANTHUS BeERLANnpiERI (Moqg.) Uline & Bray. San Juan (444).
AMARANTHUS Hysripus L. Alamo (1342).

AMARANTHUS sprnosus L. Palm Grove, Brownsville (1534).
CELOsIA PANICULATA L. Salino Station, Hidalgo Co. (1609).
CLaDOTHRIx LANUGINOSA Nutt. Boca Chica (357).

FROELICHIA CAMPESTRIS Small. Rio Grande City (1939).
FroeticH1a Drummonpit Mog. N. Hidalgo Co. (855).

PHILOXERUS VERMICULARIS (L.) R. Br. Padre Island (380).

NYCTAGINACEAE

ACLEISANTHES Greco Standley. San Juan (464).
ACLEISANTHES oBTUSA (Choisy) Standl. Roma (1386).
ALLIONIA NYCTAGINEA Michx. Clark Island (1715).
ANREDERA VESICARIA (Lam.) Gaertn. (1919).

BoerHAaviA ERECTA L. San Juan (302).

BoerHAAVIA viscosa Lag. & Rohr. Rio Grande City (1312).
Nycracinia capitata Choisy. Rio Grande City (187).
WEDELIELLA INCARNATA (L.) Cockerell. Zapata (1358).

PHYTOLACCACEAE

PHAULOTHAMNUS SPINESCENS Gray. Rio Hondo (1034).
Rivina HumitIs L. Alamo (13).

AIZOACEAE

Mo.ttvco verticittata L. Harlingen (1201).
SEesuvium verrucosum Raf. Boca Chica (1713).

CACTACEAE

ACANTHOCEREUS PENTAGONUS (L.) Britt. & Rose. Rio Hondo (1872).
ANCISTROCACTUS BREVIHAMATUS (Engelm.) Britt. & Rose. La Joya.
ANcIsTRocAcTusS ScHEERI Salm-Dyck. La Joya (1887).

AsrropHytum astrrias (Zucc.) Lemaire. Rio Grande City (1867).
CorYPHANTHA Pirtieana I. Wright. Rio Grande City (15273, Gard. No.)
CoryPHANTHA Runyonm Britt. & Rose. Rio Grande City (1875).
Do.LicoOTHELE SPHAERICA (Dietrich) Britt. & Rose. Salinefio (1876).
ECHINOCEREUS ANGUSTICEPS Clover. Linn (1870). (Type.)
Ecuinocereus Brancxi (Poselger) Palmer. Alamo (1878).
ECHINOCEREUS ENNEACANTHUs Engelm. La Joya (1874).

Ecuinocerevus Fircum Britt. & Rose. Rio Grande City (1871).
EcHINOCEREUS PAPILLOSUS Linke. Rucio (1869).

EcHINOCEREUS PENTALOPHUS (DC.) Rumpler. La Joya (1890).
EscopartaA Runyonm Britt. & Rose. Tabasco (1886).

FEROCACTUS HAMATACANTHUS (Muhlenpfordt) Britt. & Rose. Zapata

(1873).

HaAMATOCACTUS SETISPINUS Engelm. La Joya (1876).

HoMaALocePHALA TEXENSIS (Hopffer) Britt. & Rose. La Joya (1868).
LorpHopHora Wiiiiamsit (Lemaire) Coulter. Rio Grande City (187).
NEOMAMMILLARIA APPLANATA (Engelm.) Britt. & Rose. La Joya (1889).
NEOMAMMILLARIA HEMISPHAERICA (Engelm.) Britt. & Rose. La Joya

(1885).

NEoMAMMILLARIA Heypert (Muhlenpfordt) Britt. & Rose. La Joya

(1891).

NEOMAMMILLARIA MULTICEPS (Salm-Dyck.) Britt. & Rose. Raymondville

(1865).

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 91

Opuntia Atxarret Griffiths. Boca Chica.

OpunTIA ENGELMANNI Salm-Dyck. Zapata.

OpuntTIA LEPTOCAULIS DC. Pefiitas (29).

Opuntia LinpHEIMERI Engelm. Pefiitas (35).

Opuntia Scnotti Engelm. Rio Grande City (1884).

THELocActTus Bicotor (Galeotti) Britt. & Rose. Rio Grande City (1880).
Witcox1a Posexcert (Lemaire) Coulter. La Joya (1877).

PorTULACACEAE

PortTuLaca OLERACEA L. Boca Chica (352).

TALINUM ANGUSTISsIMUM (Gray) Wooton & Standl. McAllen (1001).
TALINUM REFLEXUM Cav. Tabasco (79).

TALINOPSIS FRUTESCENS Gray. Rio Grande City (481).

CARYOPHYLLACEAE
AtsInE Batpwintt Small. San Benito (617).

NYMPHAEACEAE
CAsTALIA ELEGANS Greene. Alamo (944).

RANUNCULACEAE
Crematis Drummonpi Torr. & Gray. San Juan (1).

MENISPERMACEAE
Coccutus piversiFotius DC. Santa Maria (1215).

PAPAVERACEAE
ARGEMONE ALBA Lestib. Mercedes (467).
ARGEMONE MEXICANA L. Alamo (192).
ARGEMONE PLATYCERAS Link & Otto. Chihuahua (601).

CRUCIFERAE

CAKILE AMERICANA Nutt. Boca Chica (353).

Lermium Austrinum (Millsp.) Small. La Joya (1876).

Leripium vircinicum L. Clark Island (749).

LEsQUERELLA AURICULATA (Engelm. & Gray) S. Wats. Barreda (1648).
LESQUERELLA DENSIFLORA (Gray) S. Wats. Zapata (1794).
LesQuERELLA LAstocarPA (Hook.) S. Wats. Donna (624).
NERISYRENIA CAMPORUM (Gray) Greene. San Ignacio (684).
Rapicuta Watteri (EIll.) Small. Chihuahua (597).

Rorrea NAsturtTIuM (L.) Rusby. Palm Grove, Brownsville (1795).
SeLcenta sp. Alamo (1424). Possibly a new species.

SyNTHLIPsIs BerLANDIERI Gray. Zapata (1795).

CAPPARIDACEAE

CrisTATELLA EROSA Nutt. Monte Cristo (931).
PoLANISIA TRACHYSPERMA Torr. & Gray. Rio Grande City (474).

CRASSULACEAE
LENOPHYLLUM TEXANUM (J. G. Smith) Rose. Tabasco.

ROSACEAE
Rvusvus rriviatis Michx. Brownsville (1711).

LEGUMINOSAE

Acacia AMENTACEA DC. Rio Grande City (472).
Acacta Bertanpiert Benth. Alamo (1337).

92

MADRONO | [Vol. 4

Acacia Farnestana (L.) Willd. Alamo (293).

Acacta Grecem Gray. San Juan (83).

Acacia Rormeritana Schlect. Monte Cristo (935).

DesMANTHUS DEPRESsSA (Willd.) Kuntze. San Juan (303).
CALLIANDRA ERIOPHYLLA (Benth.) Britt. Rio Grande City (1677).
BaPTisiA LEUCOPHAEA Nutt. Clark Island (769).
CassIA BAUHINIOoIES Gray. Rio Grande City (1311).
Cassta Mepscert Shafer. Edinburg (1588).

Cassa PuMILIO Gray. Zapata (679).

Crrcipium FLoRIDUM Benth. La Joya (65).
CERCIDIUM TEXANUM Gray. (1321).

CHAMAECRISTA CINEREA (Cham. & Schlecht.) Pollard. Starr Co. (1102).
CHAMAECRISTA LITTORALIS Pollard. Padre Island (1638).

Cotocanta sp. Clark Island (746). Possibly a new species.

CouRsETIA AXILLARIS Coult. & Rose. La Lomita (Mission) (1060).
TepHrosia Linpuemmert (Gray) Kuntze. Raymondville (1198).
DavBentonia Cavinittesit (S. Wats.) Standl. Boca Chica (388).
RHYNCHOSIA AMERICANA (Mill.) Vail. Clark Island (814).
RHYNCHOSIA MINIMA (L.) Medic. Weslaco (459).

ERYTHRINA HERBACEA L. Palm Grove, Brownsville (1925).
EYsENHARDTIA TEXANA Scheele. La Joya (72).

Hamosa Nutraruiana (DC.) Rydb. (754).

HoFFrMANSEGGIA CAUDATA Gray. (1275).

HoFFMANSEGGIA DENSIFLORA Benth. Boca Chica (360).

INDIGOFERA LEPTOSEPALA Nutt. Boca Chica (704).

LEUCAENA PULVERULENTA (Schlecht.) Benth. Alamo (455).

Lupinus TEXENSIS Hook. La Joya (104).

Mimosa Bertanpiert Gray. Boca Chica (393).

Mimosa srricittosa Torr. & Gray. Boca Chica (712).

PARKINSONIA ACULEATA L. Alamo (105).

ParoseLA AUREA (Nutt.) Britton. N. Starr Co. (1106).

ParoseLA HumILIs (Mill.) Rydb. Boca Chica (1242).

ParosELA NANA (Torr.) Heller. Arroyo el Tigre (694).

PAROSELA POGONATHERA (Gray) Vail. Havana (646).

PETALOSTEMON EMARGINATUS Torr. & Gray. Boca Chica (703).
PITHECOLOBIUM BREVIFOLIUM Benth. La Lomita (Mission) (1072).
PRosoPiIs JULIFLORA Var. GLANDULOSA Torr. San Juan (2).

SIDEROCARPOS FLEXICAULIS (Benth.) Small. San Juan (7).

SopHoORA SECUNDIFLORA Lag. San Benito (1743).

SorpHorA TOMENTOSA L. Boca Chica (768).

STROMBOCARPA CINERESCENS Gray. Los Fresnos (969). |
Vicia TEXANA (Torr. & Gray) Small. Rio Hondo (611). |

GERANIACEAE
GERANIUM CAROLINIANUM L. Alamo (1779).

OXALIDACEAE
TonoxaLis vioLAceA (L.) Small. Alamo (1425).
IonoxaLis sp. Palm Grove, Brownsville (1512).
MoNOoXALIS DICHONDRAEFOLIA (Gray) Small. Rio Grande City (1046).
Oxais cornicuLtaTA L. Alamo (1423).

LINACEAE

CATHARTOLINUM ALATUM Small. Boca Chica (316).
CATHARTOLINUM MULTICAULE (Hook.) Small. N. Starr Co. (1097).

ZYGOPHYLLACEAE
CovILLEA TRIDENTATA (DC.) Vail. San Ignacio (1938).
Guaracum sanctum L. Pefiitas (89A).
KALLSTROEMIA PARVIFLORA Norton. Clark Island (1308).

| 1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY

PoruieRIA ANGUSTIFOLIA Engelm. Tabasco (89).
TRIBULUS TERRESTRIS L. San Juan (299).

RUTACEAE

Amyris PARVIFOLIA Gray. Rio Hondo (242).

EsENBECKIA Runyon Morton. Santa Maria.

Hewierra parvirotia Benth. & Hook. Rio Grande City (1317).
THAMNOSMA TEXANA (Gray) Torr. Tabasco (581).
XANTHOXYLUM INSULARE Rose. Rio Hondo (1035).
XanrHoxytum Prerota H. B. K. Rio Hondo (239).

SIMARUBACEAE

‘ CasteLA Nicworsonit Hook. Rio Hondo (241).
» CCasrenaria TEXANA (Torr. & Gray) Small. Alamo (1501).

MALPIGHIACEAE

_~-CoELOSTYLIS TEXANA Torr. & Gray. Alamo (881).
Marricuia cuasra L. Alamo (553).

POLYGALACEAE

PotyeaLaA ALBA Nutt. Boca Chica (706).
Potyeata Boyxinu Nutt. Clark Island (745).
PoLYGALA MACRADENIA Gray. Rio Grande City (488).

EUPHORBIACEAE

ACALYPHA HEDERACEA Torr. La Joya (183).

ADELIA Vasey1 (Coulter) Pax & Hoff. Mission (1077).
BERNARDIA MYRICAEFOLIA (Scheele) Gray. Rio Hondo (1036).
CHAMAESYCE CoRDIFOLIA (Ell.) Small. Mission (356).
CHAMAESYCE HYPERICIFOLIA (L.) Small. Alamo (356A).
CHAMAESYCE LAREDANA (Millsp.) Small. Rio Grande City (1363).
CHAMAESYCE SERPENS (H. B. K.) Small. Rio Grande City (1364).
CROTON CILIATO-GLANDULOsUs Ort. Roma (1392).

Croton Cortestanus Kunth. Raymondville (190).

Croton ENGELMANNII Ferguson. Raymondville (1216).

Croton FRUTICULOsUs Engelm. Alamo (1483).

Croton LEUCOPHYLLUS Muell. Arg. Ojo de Agua (999).

Croron NEoMEXICANUS Muell. Arg. Edinburg (1596).

Croton PuNcTaATus Jacq. Padre Island (1635).

Croton Torreyanus Muell. Arg. Rio Grande City (1330).
Croton sp. Alamo (1776). Possibly a new species.

Diraxis HuMiLIs Engelm. & Gray. Arroyo Velefio (1786).
GALARHOEUS ARKANSANUS (Engelm. & Gray) Small. Alamo (872).
GaLaRHOEUs sp. Alamo (889). Possibly a new species.
JATROPHA Beritanpieri Torr. Rio Grande City (478).

JATROPHA SPATHULATA Muell. La Joya (39).

JATROPHA stimuLosA Michx. Santa Elena Ranch (1844).
POINSETTIA HETEROPHYLLA L. San Juan (109).

Stirtincia Torreyana Wats. Zapata (498). ie

TRAGIA NEPETAEFOLIA Cav. Palm Grove, Brownsville (1562).
TRAGIA urRTICAEFOLIA Michx. Camp Perry (1758).

CELASTRACEAE
MayTENUS PHYLLANTHOIWES Benth. Brownsville (417).
Morron1a Grecert Gray. La Joya (1791).
SCHAEFFERIA CUNEIFOLIA Gray. La Joya (73).
\( CaRDIOSPERMUM corINDUM L. Palm Grove, Brownsville (1558).
CarpiosPpeERMUM Hatricacasum L. Tabasco (205).
. + |Sarinpus Drummonni Hook. & Arn. Alamo (779).
SERJANIA BRACHYCARPA Gray. Alamo (1468).

94

MADRONO [Vol. 4

RHAMNACEAE

CoLuBRINA TEXENSIS Gray. Zapata (1033).
Conpatia opovaTa Hook. Havana (1129).

__Conpauia ostusirot1a (Hook.) Weberbauer. Rio Grande City (1371).

Karwinsk1a Humporptiana Zuce. Alamo (11).
MicroRHAMNUS ERICOIDES Gray. Rio Grande City (470).
ZiZYPHUS oOBTUSIFOLIA (Hook.) Gray. Palm Grove, Brownsville (1087).

ener)

VITACEAE

AMELOPSIS ARBOREA (L..) Rusby. Tabasco (209).
AMELOPSIS MEXICANA Rose. Barreda (1702).
Cissus incisA Desmoul. Zapata (1272).

MALVACEAE

ABUTILON AMERICANUM (L.) Sweet. Rio Grande City (185).
ABUTILON INCANUM (Link.) Sweet. Boca Chica (1238).
ABUTILON Jacauintt Don. Palm Grove, Brownsville (1505).
ABUTILON TEXENSIS Torr. & Gray. Alamo (1454).

ABUTILON TRIQUETRUM (L.) Presl. Alamo (1446).
Axsutiton Wricuti Gray. Rio Grande City (489).
CALLIRRHOE pDIGITATA Nutt. San Ignacio (685).

CIENFUEGOSIA SULPHUREA (St. Hil.) Garcke. Alamo (1259).
Gayoipes crispum (L.) Small. Roma (1391).

Hisiscus carpiopHytius Gray. La Joya (38).

_KosteLerzkya HASTATA Presl. Edinburg (1608).

Maracura uRENS Poit. Palm Grove, Brownsville (1526).

MALVASTRUM COROMANDELIANUM Garcke. Boca Chica (407).
Matvastrum spicatum (L.) Gray. Boca Chica (406).

Matvaviscus Drummonp1 Torr. & Gray. Palm Grove, Brownsville (1845).
Sma pirrusa H. B. K. Alamo (1498).

Sma Fiuires Gray. Roma (1408).

Sma wastata St. Hil. Las Comitas (1314).

Sipa panicutatTa L. Rio Hondo (1014).

Sma sprnosA L. Mission (112).

WissaDULA AMPLIssImMA (L.) R. E. Fries. Palm Grove (1564).

VIOLACEAE
CALCEOLARIA VERTICILLARIA (Ortega) Kuntze. Clark Island (723).

Gposeedoee
WaALTHERIA AMERICANA L. San Isidro Ranch (1840).

STERCULIACEAE
MeLocHIA PyrAMIDATA L. Olmito (1261).

CocHLOSPERMACEAE
Amorevuxia Wricuti Gray. Rio Grande City (486).

KOEBERLINIACEAE
KOoEBERLINIA spinosa Zucc. La Joya (64).

FLACOURTIACEAE
XYLOSMA CELASTRINUM (H. B. K.) Kuntze. Combes (1641).

TURNERACEAE
TURNERA APHRODISIACA Ward. Zapata (1686).
PassIFLoRA FOETIDA L. Mission (191).
PassiFLorA LUTEA L. Palm Grove, Brownsville (1603).

LOASACEAE
Coenen, sInuaTA Lag. Zapata (1409).

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 95

ONAGRACEAE

Gatprnsia Hartwecit (Benth.) Britton. Arroyo el Tigre (695).
GALPINSIA TUBICULA (Gray) Small. Clark Island (719).

Gavura Drummonpiu Torr. & Gray. Weslaco (623).

GavurRA vittosa Torr. Las Viboras Ranch (1100).

HarrMannia speciosa (Nutt.) Small. Pefiitas (1846).

JUSSIAEA pIFFUSA Forskal. Rio Hondo (1040).

KNEIFFIA ARENICOLA Small. Mission (169).

OrnorHerRA Drummonpit Hook. Boca Chica (351).

UMBELLIFERAE
AMMOSELINUM Poprerr Torr. & Gray. Chihuahua (573).
Arium Amoi (L.) Urban. La Joya (1895).
Eryncium compactum Small. Alamo (874).
EurRYTAENIA TEXANA (Torr. & Gray) Small. Santa Elena Ranch (1107).
HyprocoryLte UMBELLATA L. Alamo (787).

PRIMULACEAE
SAMODIA EBRACTEATA (H. B. K.) Baudo. Boca Chica (318).
SaMOLUS CUNEATUS Small. Boca Chica (1248).
Samouvus FLorisuUNDUs H. B. K. Rio Hondo (1026).

PLUMBAGINACEAE

Limontum Nasui Small. Boca Chica (355).
PLUMBAGO SCANDENS L. Alamo (785).

SAPOTACEAE

BuMELIA ANGUSTIFOLIA Nutt. Edinburg (1328).
Bumetia tyciowers Pers. La Joya (1328A).

EBENACEAE
Diospyros TexANA Scheele. Alamo (784).

OLEACEAE

ForEsTIERA ANGUsTIFOLIA Torr. Rio Hondo (1597).
‘Fraxinus Bertanpiert DC. Palm Ae ale Brownsville (1924).
MrENopoRA HETEROPHYLLA Moric. sion (1063).

GENTIANACEAE

Evstoma Russett1anum (Hook.) Griseb. Chihuahua (199).
SappaTia cAmpPEstTRis Nutt. N. Hidalgo Co. (831).
SaBBATIA CARNOSA Small. Boca Chica (702).

ASCLEPIADACEAE

METASTELMA BARBIGERUM Scheele. Palm Grove, Brownsville (1547).
PopostEMMA LONGICORNU (Benth.) Greene. San Juan (1906).
PopostemMA sp. Boca Chica (320). Possibly a new species.
VINCETOXICUM BREVICORONATUM (Rob.) Vail. E. Starr Co. (663).
VINCETOXICUM RETICULATUM (Engelm.) Heller. Barreda (1624).

CONVOLVULACEAE _
ConvotvuLus incanus Vahl. Linn (1612).
CuscuTa ARVENSIS Beyrich. Rio Hondo (230).
Cuscuta 1npEcorA Choisy. Boca Chica (390)..
Evotvutvus austnowrs L. Tabasco (40).
Irpomora carnosa Britton. Boca Chica (314).
Iromora pissecta (Jacq.) Pursh. Alamo (1457).
Ipomoea Fistutosa Mart. Alamo (294).—
Ipomora Prs-Carrar Sweet. Boca Chica (323).

96

MADRONO Av [Vol. 4

Ipomora sinuatTa Ort. Raymondville (1196).
Ipomora triripa (H. B. K.) G. Don. Tabasco (206).

PoLEMONIACEAE

Giria incisA Benth. Rio Hondo (1905).
Putiox pitosa L. var. petonsA Gray. N. Hidalgo Co. (822).

HypDROPHYLLACEAE

MariILaAUNIDIUM HIspiIpuM (Gray) Kuntze. Chihuahua (585).
MaRILAUNIDIUM JAMAICENSE (L.) Kuntze. Chihuahua (592).
MaRILAUNIDIUM MACRANTHUM (Choisy) Kuntze. Alamo (909).
PHACELIA PARVIFLORA Pursh. Combes (1760).

PHACELIA PATULIFLORA (Engelm. & Gray) Gray. Mercedes (618).

BoRAGINACEAE

CoLDENIA CANESCENS DC. Zapata (1847).

Corp1a Boissternt A. DC. Alamo (9).

EnRetTIA ELLIPTICA DC. Alamo (896).
HELIOTROPIUM CONFERTIFOLIUM Torr. Zapata (682).
HeLioTropium curassAvicum L. Boca Chica (396).
He.iorropium 1npicum L. Boca Chica (202).
HELIOTROPIUM INUNDATUM Swartz. Alamo (902).
HELIoTROPIUM PARVIFLORUM L. E. Starr Co. (659).
LirHOsPERMUM MATAMORENSE DC. Tabasco (676).
TouRNEFORTIA VoLUBILIS L. Camp Perry (1756).

VERBENACEAE

AVICENNIA NITIDA Jacq. Clark Island (1251).

CITHAREXYLUM BeriaAnpbiEeRI Robinson. Boca Chica (1694).
GoNIOSTACHYUM cITROSUM Small. Rio Hondo (1566).
HELLERANTHUS QUADRANGULATUS (Heller) Small. Chihuahua (596).
LANTANA HoRRIDA H. B. K. Pefiitas (1426).

LANTANA MACROPODA Torr. Rio Grande City (1681).

Lipp1a ALBA (Mill.) N. E. Brown. Palm Grove (962).

Lipr1a Bertannpieri Schauer. Tabasco (1676).

Liprp1a LicustRINA (Lag.) Small. Tabasco (10). !
Liprra macrostacuya (Torr.) Moldenke. Rio Grande City (1268).
PuyLa NopiFLtorA (L.) Greene. Mission (119).

VERBENA BIPINNATIFIDA Nutt. Tabasco (106).

VERBENA ci“1aTa Benth. Alamo (1450).

VersenA Hatret Small. Clark Island (736).

VERBENA OFFICINALIS L. Barreda (1695).

VERBENA xUTHA Lehm. Rio Grande City (1618).

VerRsena (hybrid?). Palm Grove, Brownsville (1519).

LABIATAE

MeELOsMON CUBENSE (L.) Small. Mission (171).

MIcroMERIA PILOsIuscULA (Benth.) Small. Camp Perry (1903).
Monarpba DispersA Small. San Juan (5).

Monarpa punctata L. Alamo (1046).

Monarpa sp. N. Hidalgo Co. (807). Possibly a new species.
SALVIA BALLOTAEFLORA Benth. La Joya (1333).

Satvia coccinea L. Raymondville (606).

Satvia Greco Gray. Brownsville.

ScuTetLar1A DrumMmonpit Benth. Mission (591).

Stacuys acrartA Cham. & Schlecht. Rio Hondo (1051).
Stacuys Drummonpi Benth. Chihuahua (602).

ZORNIA BRACTEATA (Walt.) Gmel. N. Starr Co. (1841).

Sv

1937] CLOVER: VEGETATION OF RIO GRANDE VALLEY 97

SOLANACEAE
Capsicum spaccatum L. Alamo (1113).

. CHAMAESARACHA contomwEs (Moric.) Britt. Barreda (1714).

Datura INNOxIA Safford. Roma (502).

Lycium Berianvrert Dunal. Rio Grande City (490).
LyciIuM CAROLINIANUM Walt. Olmito (1654).
Lycrium Cuateavur Standl. La Joya (74).<—
Lycium Torreyi Gray. Zapata (1301).
LYCOPERSICON CERASIFORME Dunal. Alamo (1052).
Nicotiana GLauca Graham. ‘La Joya (184).
NICOTIANA REPANDA Willd. Roma (1396).

PETUNIA PARVIFLORA Juss. Chihuahua (505).
Prysatis Carrenteri Riddell. Roma (1383).
Puysauis moniis Nutt. Clark Island (1926).
Puysauis viscosa L. Los Fresnos (1631).

PuysaLis viscosa L. var. SPATHULAEFOLIA Gray. Clark Island (728).
QuiIncuLA LoBATA (Torr.) Raf. Tabasco (598).

SoLANUM CAROLINENSE L. Alamo (1122).

SoLANUM ELAEAGNIFOLIUM Cay. Alamo (264).

SoLanum nicrum L. Alamo (917).

SoLanum rRosrratum Dunal. Roma (503).

SoLANUM TRIQUETRUM Cav. San Benito (616).

ScROPHULARIACEAE
GERARDIA HETEROPHYLLA Nutt. Boca Chica (1252).
LEUCOPHYLLUM FRUTESCENS (Berlandier) Johnston. (81).
LevucoPpHyLLuMm Frutescens f. albiflorum Clover. A JL. frutescens differt

floribus albis haud purpureo-rosaceis. Legit ad Romam, Starr Co., Texas,
Clover 492, in Herb. Mich.

LEUCcOPHYLLUM miINUs Gray. San Benito (1746).
Linaria CANADENSIS (L.) Dumont subsp. rexana (Scheele) Pennell. Mis-

sion (1787).

STEMODIA LANATA Ruiz & Pavon. Clark Island (747).

MartTyNIACEAE
Martynia FRAGRANS Lindl. Roma (120).

LENTIBULARIACEAE
Urricutaria susuLtATA L. Rio Hondo (1748).

ACANTHACEAE
Justicia Runyon Small. Brownsville (1233).
CARLOWRIGHTIA GLABRATA Fernald. Mission (1066).
RUELLIA INTERMEDIA Leonard. La Joya (118).
RUELLIA NoctiFtorA (Nees) Gray. Roma (1389).

_Rvetria tuserosa L. Tabasco (92).

SIPHONOGLOSSA DIPTERACANTHA (Nees) Heller. (1911).
TUBIFLORA ACUMINATA Small. E. Starr Co. (652).

PLANTAGIN ACEAE

PLANTAGO LANATIFOLIA (Coult. & Fish.) Small. Zapata (1922).
PLANTAGO RHODOSPERMA Decne. Clark Island (1921).
PLANTAGO virGINiIca L. var. toncirotia Gray. Tabasco (675).

RUBIACEAE
-GALIUM APARINE L. Palm Grove, Brownsville (1703).

“sMacrosipHonia MacrosipHon (Torr.) Heller. La Joya (554).

| oe

Rawnpia acuteata L. Mission (1065).
RICHARDIA BRAZILIENSIS (Moq.) Gomez. Santa Elena Ranch (1109).

98

MADRONO [Vol. 4

CUCURBITACEAE

Cucumis Ancuria L. La Joya (1897).

CucursiTa rorTipissimA H. B. K. Tabasco (1505).
IservitteaA Linpueimmeri (Gray) Greene. Alamo (1919).
MeELoTHRIA PENDULA L. Alamo (1417).

CAMPANULACEAE

Losetia BRACHYPODA A. DC. Rio Grande City (908).
Loseria Criirrort1rana L. Rio Hondo (615).
Loser1a Feayana Gray. Los Fresnos (1729).

LYTHRACEAE

AMMANNIA COCCINEA Rottb. Palm Grove, Brownsville (1523).
Her1a sariciroria Link. Havana (1126).
LyrHrum aLtatum Pursh. Mercedes (287).

COMPOSITAE

ACTINEA LINEARIFOLIA (Hook.) Greene. Zapata (692).

AcTINEA oporATA (DC.) Kuntze. Zapata (1693).

AMBLYOLEPIS SETIGERA DC. Rio Grande City (1832).

AmprosiA ELATIOR L. Mission (167).

AMBROSIA PSILOSTACHYA DC. Mission (160).

APHANOSTEPHUS RAMOsISsIMUS DC. Raymondville (1210).
APHANOSTEPHUS skirROBASIS (DC.) Trel. Clark Island (765).
APHANOSTEPHUS SKIRROBASIS var. Haru (Gray) Blake. Linn (810).

Aptopappus Drummonpi (Torr. & Gray) Blake. Los Fresnos (1732).

APLOPAPPUS PHYLLOCEPHALUS DC. Boca Chica (714).
Arctium Lappa L. Roma (1940).

Aster Exitis Ell. San Juan (441).

Aster Patmert Gray. Rio Grande City (1380).

ASTER sPINosus Benth. Pharr (789).

BaccHaris GLuTINosA Pers. Boca Chica (399).

BaccHaRis TEXANA (Torr. & Gray) Gray. Clark Island (1726).
BAHIA ABSINTHIFOLIA Benth. Rio Grande City (1359).
BorrRIcHIA FRUTESCENS (L.) DC. Clark Island (330).
CALYPTROCARPUS VIALIS Less. Mission (170).

CENTAUREA AMERICANA Nutt. Palm Grove, Brownsville (951).
Cirstum UNDULATUM Nutt. ? San Benito (627).

CLAPPIA SUAEDAEFOLIA Gray. Mercedes (619).

Conyza Couttert Gray. Camp Perry (1757).

CorREOPSIS CARDAMINIFOLIA (DC.) Torr. & Gray. Clark Island (571).
Dyssop1a Bertanpiert (DC.) Blake. Zapata (1827).
DyssopIA TEPHROLEUCA Blake. Rio Grande City (1825). Type.
Ecuipta ALBA (L.) Hassk. Alamo (873).

ERIGERON CANADENSIS L. Havana (1131).

ERIGERON REPENS Gray. Boca Chica (713).

Ericeron TENUIS Torr. & Gray. Palm Grove (1834).
EUPATORIUM AGERATIFOLIUM DC. Barreda (1647).
Evparorium azurEum DC. Barreda (1740).

EvupatTorium oporatum L. Palm Grove, Brownsville (1835).
Evax mutricautis DC. E. Zapata Co. (1864).

FLorEsTINA TRIPTERIS DC. San Juan (468).

FRANSERIA CONFERTIFLORA (DC.) Rydb. Zapata (690).
GAILLARDIA PULCHELLA Foug. Chihuahua (193).

GNAPHALIUM SPATHULATUM Lam. Rio Hondo (1044).
GocHNATIA HYPOLEUCA (DC.) Gray. San Ignacio (934).
GRINDELIA COLEPsSIS Blake. Brownsville.

GUTIERREZIA ERIOCARPA Gray. Boca Chica (389).

HELENIUM LINIFOLIUM Rydb. Raymondville (1769).
HeELENIUM MICROCEPHALUM DC. Mission (163).

HELENIUM QUADRIDENTATUM Labill. Alamo (906).

oF,

1937 | CLOVER: VEGETATION OF RIO GRANDE VALLEY 99

HELIANTHUS ANNUUsS L. Zapata (1828).

HELIANTHUS CucUMERIFOLIUs Torr. & Gray. Raymondville (1200).
HETEROTHECA SUBAXILLARIS (Lam.) Britt. & Rusby. Combes (1765).
Iva ancustirotia Nutt. Boca Chica (328).

Iva cm1ata Willd. Tabasco (213).

LepacHys cotumnaris (Sims) Torr. & Gray var. puLCHERRIMA (D. Don)

Torr. & Gray. Mission (110).

10.

LycopEesmM1A TEXANA (Torr. & Gray) Greene. N. Hidalgo Co. (824).
MELAMPODIUM CINEREUM DC. HE. Starr Co. (653).

OTHAKE ROBUSTUM Rydb. Clark Island (733).

OTHAKE TEXANUM (Hook.) Bush. Tabasco (93).

ParTHENIUM HysteropHorus L. Mission (114).

PEREZIA RUNCINATA Lag. Barreda (1622).

PERITYLE MicroGLossa Benth. Rio Hondo (1742).

PiucHea CAMPHORATA (L.) DC. Boca Chica (367).
PSILOSTROPHE GNAPHALODES DC. San Ignacio (1689).
Rupseckia Bicotor Nutt. San Manuel Ranch (809).

SELLOA GLUTINOSA Spreng. Los Fresnos (1731B).

SENECIO AMPULLACEUS Hook. Palm Grove, Brownsville (1833).
SENECIO GLABELLUS Poir. Brownsville (1644).

Srmms1a cALvA (Gray & Engelm.) Gray. Rio Grande City (1355).
PYRRHOPAPPUS GRANDIFLORA (DC.) Greene. San Benito (1928).
PyRRHOPAPPUS MULTICAULIS (DC.) Greene. Port Isabel (1649).
Soncuus oLeRAceus L. San Juan (445).

Tricuocoronis Wricutiu (Torr. & Gray) Gray. Alamo (876).
Trixis RADIALIS (L.) Kuntze. Alamo (1428).

VARILLA TEXANA Gray. Roma (1351).

VIGUIERA STENOLOBA Blake. Mission (173).

XANTHIUM sPECcIOSUM Kearney. Roma (500).

VERBESINA ENCELIOWES (Cov.) Benth. & Hook. Tabasco (172).
VERBESINA VIRGINICA L. Olmito (1645).

ZEXMENIA BREVIFOLIA Gray. Tabasco (90).

ZEXMENIA HIspipA (H. B. K.) Gray. Rio Grande City (1673).
ZINNIA PUMILA Gray. Rio Grande City (479).

University of Michigan,
Ann Arbor, January 1, 1936.

LITERATURE CITED

Baitry, Vernon. Biological survey of Texas. U.S. Dept. Agric., Biol.
Surv., N. Am. Fauna No. 25. 1905.

Barritett, H. H. A method of procedure for field work in tropical Ameri-
can phytogeography based upon a botanical reconnaissance in parts
of British Honduras and the Petén forest of Guatemala. Carnegie
Inst. Washington, Publ. No. 461: 1-25. 1935.

Becx, M. W., and B. H. Henpricxson. Soil survey of Cameron County,
Texas. U.S. Dept. Agric., Bur. of Chem. and Soils, No. 17, series
1923.

Berianpier, L. Espedicion scientifica del general Terdn 4 Tejas. Boletin
de la Sociedad Mexicana de Geografia y Estadistica 5: 125-133. 1857.

Bray, Wm. L. The ecological relations of the vegetation of western
Texas. Bot. Gaz. 32: 99-123, 195-217, 262-291. 1901.

The mistletoe pest in the southwest. U. S. Dept. Agric.,
Bur. Plant Industry, Bull. No. 166. 1910.

Britron, N. L., and J. N. Rost. The Cactaceae. Carnegie Inst. of Wash-
ington, Publ. No. 248. 1919-1923.

Crements, F. L. Plant succession—an analysis of the development of
vegetation. Carnegie Inst. of Washington, Publ. No. 242. 1916.
Crover, E. U. Astrophytum asterias in the United States. Rhodora 37:

227-228. 1932.
——_———. Hchinocereus angusticeps, a new species from the Lower
Rio Grande Valley, Texas. Rhodora 37: 77-80. 1935.

100

11.
12.

13.

14.
15.

16.
17.

18.

Wh

20.

MADRONO [Vol. 4

Coox, O. F. Change of vegetation on the south Texas prairies. U. S.
Dept. Agric., Bur. Plant Industry, Cire. No. 14. 1908.

Coorer, W. S. The broad-sclerophyll vegetation of California. An eco-
logical study of the chaparral and its related communities. Carnegie
Inst. of Washington, Publ. No. 319. 1923.

Courter, J. M. Upon a collection of plants made by Mr. G. C. Nealley,
in the region of the Rio Grande, in Texas, from Brazos Santiago to
El Paso County. Contr. U.S. Nat. Herb. 1: 29-61. 1890.

. Botany of western Texas. Contr. U. S. Nat. Herb. 2:
1-588. 1891-1894.

Cowtrs, H. C. Review of “Plant Succession” by F. E. Clements. Bot.
Gaz. 68: 477-478. 1919.

Druper, O. Handbuch der Pflanzengeographie. Stuttgart. 1890.

Emory, W. H. Report on the United States and Mexican Boundary Sur-
vey 1(1): xvit258. 1857.

ENGELMANN, G., and A. Gray. Plantae Lindheimerianae. Boston Jour.
Nat. Hist. 5: 210-264. 1845.

ENGELMANN, G. Cactaceae of the Boundary. In Emory, Wm. H., Report
on the United States and Mexican Boundary Survey 2(1): 1-78, pls.
1-75. 1858.

Enectier, A. Die Pflanzengeographische Gliederung Nord-Amerikas. Ab-
druck, aus dem Notizblatt des Konigl. Bot. Gart. Berlin, Appendix
IX. 1902.

Foscur, E. J. Physiography of the Lower Rio Grande Valley. Pan-
American Geologist 57: 263-267. 1932.

Gray, A. Plantae Lindheimerianae. Boston Jour. Nat. Hist. 6: 141-240.
1850.

HarsHsBercer, J. W. Phytogeographic survey of North America. In
Engler und Drude, Die Vegetation der Erde 13: vi+789. 1911.

Havarp, V. The Mezquit. Am. Nat. 18: 451-459. 1884.

—_—_—_——. Report on the flora of western and southern Texas. Proc.
U. S. Nat. Mus. 8: 449-533. 1885.

Hooxer, W. J. Notice concerning Mr. Drummond’s collections made
chiefly in the southern and western parts of the United States. Com-
panion to Bot. Mag. 1: 39-49, 95-101, 170-177. 1835; 2: 60-64. 1836.

Lunpet1, C. L. Preliminary sketch of the phytogeography of the Yucatan
Peninsula. Carnegie Inst. of Washington, Publ. No. 436. 1934.

. Merriam, C. H. Life zones and crop zones of the United States. U. S.

Dept. Agric., Biol. Surv., Bull. No. 10. 1898.
Scuotr, A. V., and C. C. Parry. In Emory, Wm. H., Report on the
United States and Mexican Boundary Survey 1(2): viiit+174. 1857.

. Scuuuz, E. D., and R. Runyon. Texas cacti; a popular and scientific

account of the cacti native of Texas. Trans. Texas Acad. Sci. includ-
ing Proc. 14: 1-181. 1930.

Sma, J. K. The palmetto-palm, Sabal texana. Jour. N. Y. Bot. Gard.
28: 132-143. 1927.

SpautpinGc, M. V. Biological relations of certain desert shrubs. Bot. Gaz.
38: 122-138. 1904.

. Tarp, B. C. Structure of Texas vegetation east of the 98th meridian.

Univ. Texas Bull. No. 2602. 1926.

. Torrey, J. Botany of the Boundary. In Emory, Wm. H., Report on the

United States and Mexican Boundary Survey 2(1): 27-270, pls. 1-61.
1858.

. Trowsrincr, A. C. Tertiary and quaternary geology of the Lower Rio

Grande Valley region, Texas. U.S. Geol. Surv. Bull. No. 837. 1932.

. Warmine, E. U. G. Occology of plants. Oxford, Clarendon Press. 1909.
. Weaver, J. E., and F. R. Crements. Plant ecology. New York. 1929.
. Winker, C. H. The botany of Texas: an account of the botanical in-

vestigations in Texas and adjoining territory. Univ. Texas Bull. No.
183 L915:

1937] _ NOTES AND NEWS 101

NOTES AND NEWS

On June 30, 1937, Dr. Willis Linn Jepson, Professor of Bot-
any, University of California, retires from active service. An
alumnus of the institution, he joined the faculty in 1894 and has
since devoted most of his attention to the study of the California
flora. Among his many well known publications on this subject
the following may be mentioned: “The Silva of California,”
1911; “‘The Trees of California,” 1909; “A Flora of California,”
1909-1987 (Volumes 1 and 2 completed) ; “A Manual of Flower-
ing Plants of California,” 1925. Dr. Jepson directed the organi-
zation of the California Botanical Society in 1913 and was for
many years its president. He founded “MaproNo” in 1916 and
acted as editor of the journal until 1934.

Dr. Lincoln Constance has recently been appointed Assistant
Professor of Botany and Assistant Curator of the Herbarium at
the University of California, Berkeley. Since 1933 he has held
a similar position at Washington State College, Pullman. A
graduate of the University of Oregon, Dr. Constance entered the
University of California as teaching assistant in botany, and in
19383 received from that institution the degree of Doctor of

Phliosophy.

The publications listed below are of special interest to botan-
ists of western North America.

Recent issues of the University of California Publications in
Botany: “A systematic study of the genus Eriophyllum,” by Lin-
coln Constance (Univ. Calif. Publ. Bot. 18: 69-136, pls. 3-8, text
figs. 16, 1987. $.50); “An unusual ascomycete in the shells of
marine animals [Didymella conchae Bonar|,’ by Lee Bonar
(Univ. Calif. Publ. Bot. 19: 187-194, pl. 22, text fig. 1, 19386.
$.25) ; ‘“Iridophycus, with special reference to the South American
species,’ by William Albert Setchell and Nathaniel Lyon Gardner
(Univ. Calif. Publ. Bot. 19: 195-244, pls. 23-29, 1987. $.50).

“Moisture relations in the chaparral of the Santa Monica
Mountains, California,’ by Harry Lloyd Bauer (Ecological Mono-
graphs 6: 409-454, July, 1936).

Publications on the Templeton Crocker expedition of the
California Academy of Sciences, 1982: No. 31, “A preliminary
report on the algae,” by William Albert Setchell (Proc. Cal. Acad.
Sci. ser. 4, 22: 65-98, pls. 83-25, text fig. 1, 1987); No. 82, ““The
plant genus Coldenia in the Galapagos Islands’? by John Thomas
Howell (Proc. Calif. Acad. Sci. ser. 4, vol. 22: 99-110, pls. 26, 27,
1937).

“A key to the grasses of Montana based upon vegetative char-
acters,’ by C. Leo Hitchcock (Published by the author, University
of Montana, Missoula, Montana, pp. 1-28, pls. 1-8, 1986. $.50).

102 MADRONO [Vol. 4

This key, of convenient size and based upon characters observable
at any time during the growing season, requires as aids only a
millimeter rule and a lens of ten power magnification. It should
thus be of great service in field identification of grasses. The
plates comprise 190 excellent figures illustrating for the most
part ligule characters of nearly as many species of grasses.

California Forest and Range Experiment Station: Staff Publi-
cations, January 1, 1916, to April 30, 1987. This bibliography
includes publications of forty-three authors and lists approxi-
mately two hundred titles of important papers on forestry and
related subjects (Berkeley, California: compiled by A. M.
Avakian, Librarian. Mimeographed).

“Leaflets of Western Botany,’ owned and published by Alice
Eastwood and John Thomas Howell of the California Academy of
Sciences, San Francisco, has completed its first volume. The
volume contains twenty numbers issued from January 16, 1932 to
November 12, 1986, and consists of over one hundred articles on
various phases of western botany.

Volume I of “Contributions from the Dudley Herbarium,”
1927-19386, closed with number 7, “A resurrection and revision of
the genus Iliamna Greene” [| Malvaceae] by Ira L. Wiggins (Stan-
ford University Press, pp. 213-229, pl. 20). With this number
appears also the volume index and title-page. The volume consists
of twelve articles on the taxonomy and distribution of western
plants by the following authors: L. R. Abrams, Elmer Applegate,
Rimo Bacigalupi, Roxana S. Ferris, Howard E. McMinn, and
Ira L. Wiggins.

On July 14, 1987, Mr. Bayne Beauchamp of Honolulu, Mr.
Charles Belshaw of Oakland, student at the University of Cali-
fornia, and Mr. Ole Olson of Tri-State University, Indiana, sailed
on the Japanese liner Heityo Maru for South America with the
objective of making botanical collections for their respective in-
stitutions. Their first objective is the Inter-Andean Valley of
Peru in which region they will collect for some weeks. In Sep-
tember Mr. Beauchamp will return to Berkeley while Mr. Bel-
shaw will continue collecting for several months southward along
the Andes. Mr. Olson will cross the cordilleras into the rain
forest of the upper Andes and will then travel by boat down the
Amazon to the Atlantic Coast. He will return to Tri-State Uni-
versity by way of Rio de Janeiro or Para. The collectors are
carrying equipment for taking color photographs.

Dr. C. Leo Hitchcock of the Department of Botany of the
University of Montana at Missoula has accepted a position as
Assistant Professor of Botany at the University of Washington,
Seattle. |

1937] PROCEEDINGS OF CALIFORNIA BOTANICAL SOCIETY — 103

During June, 1937, Dr. Theodor Just of the Department of

Botany, University of Notre Dame, Indiana, visited botanical

institutions of the San Francisco Bay region in the interests of
the ““American Midland Naturalist’’ of which he is editor.

Dr. Ira L. Wiggins of the Department of Botany, Stanford
University, California, is spending a part of this summer at the
Royal Botanic Gardens at Kew, consulting type specimens in con-
nection with his researches on the flora of the Sonoran Desert of
northern Mexico.

“Wild Flowers of Southern California’? by Carl Thurston is
described on the title page as “‘an easy key to the names of
flowers, ferns, and trees, with 547 illustrations from photographs
taken by the author.” By attempting to cover only a limited
geographic area, this book is more complete than the usual popu-
lar handbook. It is designed on an original plan as a field index,
the purpose of which is to supply quickly the names of plants
with technical terms, keys, and descriptions reduced to a mini-
mum. Two hundred and fifty-two tables comprise the bulk of
the volume. These are supplemented by a short list of defini-
tions of botanical terms, a list of families and genera, and an
index of the scientific and common names (Esto Publishing Com-
pany, Pasadena. 412 pp. 1936. $4.00).

PROCEEDINGS OF THE CALIFORNIA
BOTANICAL SOCIETY

Thursday, April 1, 1987. A meeting was held at 8:00 p. m.
in Room 2098, Life Sciences Building, University of California,
Berkeley. Lecture: ‘‘Farthest South America” by Mrs. Ynes
Mexia. Mrs. Mexia recounted her experiences during a recent
botanical collecting trip to Tierra del Fuego.

Thursday, April 22, 1937. A meeting was held at 8:00 p. m.
in Room 460, Physiology Building, Stanford University. Lecture:
“In Africa and Australia after Insectivorous Plants’ by Francis E.
Lloyd, Professor of Botany, Emeritus, McGill University. The
lecture was illustrated by slides showing photographs of habitats
of Utricularia and other insectivorous plants and microphoto-
graphs, drawings, and charts of the insect trapping mechanisms.

At a business meeting of the recently organized Santa Barbara
Branch of the California Botanical Society on April 14, 1937, the
following offieers were elected: President, Mr. Maunsell Van
Rensselaer; Vice-President, Mrs. Hugh Dearing; Secretary-
Treasurer, Miss Ruth Hartwell. On the evening of March 10,
1937, at the Santa Barbara Museum of Natural History, Professor
H. E. McMinn of Mills College, California, addressed the Branch

on “The evolution of trees as represented by the trees of Santa

104 MADRONO [Vol. 4

Barbara.’ On Sunday, May 2, 1937, a field trip was taken to
Figueroa Mountain. About fifty persons attended. Leaders
were: Hugh Dearing, Ruth Hartwell, Irma Cooke, Father
Seraphin Muller, and M. Van Rensselaer. The Santa Barbara
Branch of the Society now numbers sixty-five members: Mrs.
Dorothy Crofton Atkins, Mrs. Alexander Baring, Mr. Antonio
Beatrice, Mrs. Robert Woods Bliss, Mr. William Body, Miss
Ellen Chamberlain, Mrs. Philip S. Chancellor, Miss Pearl Chase,
Miss Helen Clarke, Dr. Frederic E. Clements, Mrs. George Clyde,
Mrs. Olivia L. Converse, Mrs. E. A. Danielson, Mrs. Hugh Dear-
ing, Mr. William R. Dickinson, Mrs. Malcolm Douglas, Mr. Wil-
liam North Duane, Mr. Frank E. Dunne, Mrs. George S. Edwards,
Mrs. Muriel Edwards, Mrs. G. Hillyer Garvin, Mrs. E. Palmer
Gavit, Dr. Adele Lewis Grant, Mrs. Clifford Greene, Mrs. Clin-
ton B. Hale, Mrs. Girard van B. Hale, Mrs. William Hart, Miss
Caroline Hazard, Mr. Bernard Hoffman, Mrs. Frank Holtzbauer
(Chairman, Native Daughters of the Golden West), Mr. W. M.
James, Mrs. Kirk B. Johnson, Mr. Bert Kallman, Mr. Frederick B.
Kellam, Mrs. W. F. Kelly, Mrs. Lora J. Knight, General William
Lassiter, Miss Lorraine Lawton (Secretary, San Foca Garden
Guild), Mrs. J. D. Lewis, Mrs. Francis V. Lloyd, Dr. Frances Long,
Mrs. August C. Magnus, Mr. Emmett Martin, Mrs. Henry P.
Moseley, Father Seraphin Muller, Mrs. Charles Nelson, Mrs.
William P. Nelson, Mrs. George M. Newell, Dr. Asbjorn P.
Ousdal, Mrs. DeWitt Parshall, Mr. Reginald C. Robbins, Mr. C.
S. Robinson, Mr. E. D. Rowe, Mr. Hobart O. Skofield, Mrs. Otis
S. Southworth, Mr. George L. Stebbins, Mrs. George F. Steed-
~man, Mrs. Anne Stow-Fithian, Mr. M. Van Rensselaer, Mr. L. D.
Waller, Margaret Ely Webb, Dr. Harrington Wells, Mrs. George
G. Whitelaw, Mr. Harry S. Wilson, Mr. W. H. Yule.

The Santa Barbara Branch of the California Botanical Soci-
ety is governed by a council made up of a representative from
each of the following organizations or institutions: Blaksley
Botanic Garden, Santa Barbara Museum of Natural History,
Garden Club of Santa Barbara and Montecito, Little Garden
Club of Santa Barbara, Santa Barbara County Horticultural Soci-

ety, The Santa Barbara Gardener, Flower Seed Industry of |

Santa Barbara County, United States Forest Service, Santa Bar-
bara County Forest Department, Santa Barbara City School Sys-
tem, Santa Barbara County School System, Santa Barbara State
College, Santa Barbara School of Nature Study, St. Anthony’s
College, Plans and Planting Committee, Community Arts Associ-
ation of Santa Barbara, Carnegie Institution of Washington.

A meeting of the Santa Barbara Branch was held at the Santa
Barbara Museum of Natural History, Friday, June 18th, at 8: 00
p.m. Mr. H. M. Butterfield of the College of Agriculture, Unig
versity of California, Berkeley, spoke on “Early Introductions of |
Ornamental Plants to Santa Barbara and California.” |

Complete Your Files!

MADRONO.

A West American Journal of Botany

“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges.

Volume I, 1916-1929. . . $5.00
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Single numbers ....<.. O19

There is only a limited supply of Vol-
umes [| and II.

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and orders to:

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Stanford University, California

- VOLUME IV NUMBER 4

MADRONO

A WEST AMERICAN JOURNAL OF
BOTANY

we

Contents

_ A New Tuaticrrum rrom Mount Rainier, WasHinoton, Harold St. John 114

5 EDLINGS FROM POLYEMBRYONIC SEEDS OF Evucenta Hooxeri, Arthur M.
LEC Ni, (GJ RBIS IES AE CS a a a Ten ee aOR N en RE cee 115

SEMEL YZ Ma AG ONES SGT GSA Ue a On, SE i OO ye MS ae pe 119

) Atea Pryromornuta, Herbert F. Copeland ................ 200s eee 120
Naruantet Lyon Garpner, William A. Setchell .......... 40 slab 1 an aree aoe a 126

‘w CALIFORNIAN Species OF Bropiaga, Robert F. Hoover .............. 130

EW: St. John, Flora of southeastern Washington and of adjacent Idaho
See ATMO ONSEANCE Ye army Ml Oy lio Ua dias) 132

Published at Lime and Green Streets, Lancaster,
Pennsylvania

October, 1937

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asrams, Stanford University, California.

Dr. Lincotn Constance, University of California, Berkeley.

Dr. H. F. Coretann, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Dr. P. A. Munz, Pomono College, Claremont, California.

Business Manager—Dr. Davm D. Krecx
Lime and Green Sts., Lancaster, Pennsylvania
or
Carnegie Institution of Washington
Stanford University, California

Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

Established 1916. Published quarterly. Subscription Price $2.50 per year.
Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17%,
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bers $0.75.

Papers up to 15 or 20 pages are acceptable. Longer contributions may
be accepted if the excess costs of printing and illustration are borne by the
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Published at Lime and Green Streets, Lancaster,
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CALIFORNIA BOTANICAL SOCIETY

President: Dr. F. W. Foxworthy, Berkeley, California. First Vice-Presi-
dent: Dr. Ira L. Wiggins, Department of Botany, Stanford University. Sec-
ond Vice-President: Dr. Philip A. Munz, Department of Botany, Pomona
College, Claremont, California. Treasurer: Dr. David D. Keck, Carnegie In-
stitution of Washington, Stanford University. Secretary: Miss Ethel Crum,
4C04 Life Sciences Building, University of California, Berkeley.

Annual membership dues of the California Botanical Society are $2.00,
$1.50 of which is for a year’s subscription to Madrofio. Dues should be
remitted to the Treasurer. General correspondence and applications for
membership should be addressed to the Secretary.

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1937] SHREVE: VEGETATION OF BAJA CALIFORNIA 105

THE VEGETATION OF THE CAPE REGION OF BAJA
CALIFORNIA

Forrest SHREVE

The southern end of the peninsula of Baja California is vir-
tually an island from the biological standpoint on account of its
effective isolation from the nearest areas of similar climate. It
is separated from the Mexican mainland by the Gulf of Cali-
fornia, with an average width of 150 kilometers (93 miles), and
from the mesic highlands of northern Baja California by an arid
stretch of 550 kilometers (342 miles).

The groundwork of the vegetation of the peninsula is desert.
In the north there are two large mountain ranges, the Sierra
Juarez and the Sierra San Pedro Martir, with summits clothed
by coniferous forest. Between them and the Pacific lies an area
of chaparral, desert-chaparral transition and some grassland.
Between the mountains and the Gulf of California is the narrow
band of desert which connects the continental and peninsular
parts of the Sonoran Desert. In the extreme south is another
series of mountains, the Sierra Giganta, Sierra Laguna and
Sierra Victoria, which reach altitudes of 1800 to 2400 meters
(5900 to 7875 feet) and have an area of about 830 square kilo-
meters (300 square miles) above an elevation of 1000 meters
(3280 feet). Outside the Viscaifio Desert and the Magdalena
Plain the entire surface of the peninsula is rugged or mountain-
ous, and there are a few scattered ranges with small areas above
1000 meters (see map, fig. 1).

The northern mountains and the chaparral region are very
regularly visited by winter rains. The southern mountains
usually have copious summer rains. In the intervening area,
latitude 26° to 30° N., the rainfall is low and sometimes there
is none for three or four consecutive years.

The botanist who traverses the peninsula from north to
south is impressed by the steady loss of familar plants and the
constant appearance of new species throughout the entire dis-
tance of 1300 kilometers (800 miles). South of latitude 30° N.,
in the desert part of the peninsula, there is little change in the
physiognomy of the vegetation in spite of the gradual changes
in the flora. South of Comondi, at latitude 26° N., the types of
vegetation gradually become more numerous and the communi-
ties found in different situations and habitats become more un-
like each other than is the case in the central region. With the
enriched vegetation comes a more rapid change in the flora.

The distinctive character of the fauna and flora of the “Cape
Region,” or “Cape District,’ has been shown by Bryant (1),
Brandegee (4), Nelson (6), Grinnell (5) and others. Nelson

MaproXxo, Vol. 4, pp. 105-136, October 4, 1937.

ocT 4 1937

106 MADRONO [Vol. 4

has mapped the life zones of Baja California (6, pl. 82), extend-

ing the arid tropical zone northward from the cape to latitude
27° 30’ N., and comprising in the lower sonoran zone the north-
ern part of the central desert region and also the higher eleva-
tions of the southern mountains. Grinnell has studied the dis-
tribution of the birds of Baja California and has demonstrated
strong differentiation centers in the high mountains of the north
and of the Cape Region south of La Paz, as well as weaker
centers at latitude 27° N. and on the Pacific islands. Brandegee,
working over forty-five years ago, made the most thorough
botanical exploration of the Cape Region that has yet been car-
ried out and published three papers (2, 3, 4) which must long
serve as the foundation for further work in southern Baja
California.

During a visit to the southern part of the peninsula in the
spring of 1935 I was interested in locating the southern limit of
desert plants and desert types of plant communities, in determin-
ing the character of the vegetation of the Cape Region, and in
comparing the southern edge of the desert with its termination
in southern Sonora (7).

South of La Paz the topography of the peninsula is dominated
by the Sierra Laguna and Sierra Victoria, which together form a
narrow range with seven sharp peaks, steep sides, and outwash
slopes which fall at sharp gradients. These mountains are
wholly granitic, and the coarse angular character of their eroded
material is responsible for the steepness of their detrital slopes
and for the broad sandy streamways which pitch down to the
sea.
Northward from La Paz the topographic pattern of Baja
California is relatively simple for about 250 kilometers (155
miles). In this stretch lies the Sierra Giganta, which hugs the
Gulf coast with an unbroken ridge from 500 to 1000 meters
(1640 to 8280 feet) in altitude, and culminates in a rounded
peak of 1766 meters (5775 feet) at latitude 26° 7’ N. The east-
ern face of the Sierra Giganta is very precipitous, while the
western slope falls gradually to the Magdalena Plain. The es-
carpment which faces the Gulf is genetically related to the simi-
lar ones found on the eastern side of the Sierra Nevada, San
Jacinto, Cuyamaca, Juarez and San Pedro Martir ranges. With
rapidly falling elevation the escarpment crosses the peninsula
west of La Paz and runs into the Pacific coast a short distance
north of Todos Santos. The numerous drainageways on the
west slopes of the Sierra Giganta are responsible for the build-
ing of the Magdalena Plain. At present, in a stretch of 200 kilo-
meters there are only five large canyons which discharge their
flood waters to the Pacific or the coastal lagoons. The plain is
covered with thousands of small playas or dry lakes, which
testify to the lack of established drainage. The rock material
of the Sierra Giganta is very largely volcanic. Also there are

107

VEGETATION OF BAJA CALIFORNIA

SHREVE

1937 |

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108 MADRONO [Vol. 4

voleanic hills immediately east of La Paz and in an irregular
series running southward along the Gulf coast, broken between
Bahia de las Palmas and Punta Arena del Sur, and terminating
in a large area in and around the Sierra de la Trinidad, north-
east of San Jose del Cabo.

Southern Baja California, in contradiction to the northern
section, receives more rain on the east coast than on the west.
There is occasional winter rain west of the mountains, but the
summer rains which visit the east coast and the mountains are
much more copious and certain. Summer rain also occurs in the
Sierra Giganta, in the Sierra de Zacatecas, west of Concepcion
Bay, and extends sporadically northward to the mountains of
California.

The Cape Region not only has a higher rainfall than the
central section of the peninsula but receives it in the most favor-
able season. To these facts must be attributed the termination
or localization of desert and the existence in the Cape Region
of vegetation with a higher water requirement. The plant life
of the region has developed under conditions of both geographic
and climatic isolation. The available geological evidence indi-
cates that these isolating features have undergone little change
during most of the period in which the angiosperms have domi-
nated the vegetation of the earth. It would, therefore, be a
matter of considerable interest to compare the flora and vegeta-
tion of this ancient area with the region of summer rainfall in
Sonora and Sinaloa, with the long stretch of desert which it
terminates, and with the nearest mesic areas in Baja California.
Such a discussion of the relationships of the flora will have to
await a much more thorough exploration and study of the areas
involved.

Brandegee (4) tabulated by families the flora of the Cape
Region as known at that time, finding a total of 732 flowering
plants and ferns, of which number 146 are confined to the high
mountains and 586 to the lowlands. He found that 362 species
are common to central and northern Baja California and 494
common to the mainland of Mexico. Out of 390 genera there
are 230 which are represented by a single species, indicating a
ratio of genera to species similar to that found in island floras.
There were four genera and 72 species which appeared to be
endemic to the Cape Region as far as known in 1892. An im-
portant phase of the floral relationships which Brandegee was
not able to touch concerns the number of species on the nearest
analagous part of the Mexican mainland which are absent from
the Cape District. It is probable that this number is more than
twice the total given by Brandegee for the Cape flora.

Owing to the character of the topography there are no ex-
tensive areas in the Cape Region with a relatively uniform set
of physical conditions. The largest are the plain of La Paz,
which lies between that town and the Sierra Laguna, the rolling

1937] SHREVE: VEGETATION OF BAJA CALIFORNIA 109

GTUCSON

VEGETATION OF
BAJA CALIFORNIA

MOUNTAIN FOREST
ABOVE tOOOM.

CHAPARRAL AND
TRANSITION

FOREST OF THE
CAPE REGION

KILOMETERS
L_- me — ma — ms — oe —

°

Fig. 1. Vegetation of Baja California.

limestone ridges west of the plain of La Paz, and part of the
outwash slope which falls from the eastern base of the Sierra
Victoria down to the Bahia de las Palmas. These three areas
are representative of the lowland vegetation of the Cape Region.
On the Pacific coast south of Todos Santos the vegetation is of
a much more xeric stamp. Above 1000 meters the vegetation of
the mountains is of a more mesic character.

The bajadas and plains of the Cape Region are either purely

110 MADRONO [Vol. 4

granitic or else of purely volcanic material, the former predom-
inating. The granite soil is a light gray loam which is largely
covered by sand in the plain of La Paz and is coarser and more
homogeneous on the bajadas immediately east and west of the
mountains. There is a poor development of small drainage-
ways, due to the porosity of the soil and the high mobility of the
surface. The streams originating high in the mountains have
cut deep and abrupt channels midway in their course across the
bajadas, and their floods move over a bed with very coarse white
sand and remarkably few stones and boulders. The gradient of
these arroyos is commonly as great as 1:10, and their courses
are very direct, free of meanders, islands and relicts of former
levels.

The voleanic areas south of La Paz are irregular in their
topography and mainly covered by shallow clay soils. Their
vegetation below 500 meters is desert and between 500 and 1000
meters is different only in a scattering representation of species
from the Cape forest and in a slightly greater density of stand.

The Cape forest below 1000 meters is distinctly xeric. Its
height ranges from 6 to 14 meters (19 to 45 feet), and it varies
greatly in density, composition and the growth forms which are
represented. Certain areas are dominated by slender legumi-
nous trees and others by stout-stemmed trees with low spread-
ing branches. The canopy of the forest is usually open and
always extremely irregular. It is rarely that a single species of
tree forms as much as 80 per cent of the stand, except in the
case of Jatropha cinerea. The low interlacing branches of this
tree are an obstacle to progress through the forest. In fact an
open floor is found only in glades along arroyos where Lysiloma
candida, L. microphylla and Cercidium peninsulare are dominant.
Cacti are almost omnipresent, Pachycereus pecten-aboriginum
being most abundant where the trees are thickest, and Lemaireo-
cereus Thurberi, Machaerocereus gummosus and Opuntia cholla most
common in the open situations. Shrubs are almost invariably
abundant and in slightly moist situations contribute to the forma-
tion of impenetrable thickets. One of the commonest shrubs is
Tecoma stans, which has height and stoutness of stem which
almost give it the rank of a tree. Several composite shrubs
equal the trees in height and their flowers may be seen projecting
from the tallest limbs. The commonest of these are Viguiera
tomentosa, V. deltoidea, Alvordia fruticosa and Eupatorium sagit-
tatum. The polygonaceous vine Antigonon leptopus is abundant
in all but the driest situations and its clusters of brilliant crimson
flowers do much to give vivid color to a floral display in which
yellow is predominant.

The relatively rich composition of the Cape forest, the close
mingling of trees of different height and branching habit, the oc-
currence of erect compact shrubs, broad poorly branched ones
and semi-scandent ones, of cacti, yuccas and vines, gives much
of the Cape Region the air of an impoverished tropical jungle.

1937 ] SHREVE: VEGETATION OF BAJA CALIFORNIA 111

The vegetation of the Cape Region bears some resemblance
to the thorn-forest of Sinaloa (8) in height, density, and many
of the growth forms which are to be found. Nowhere is the
Cape forest dominated by the thorny acacia type of tree, as is
the case in Sinaloa, and it is scarcely appropriate to designate it
as ‘“‘thorn-forest.”

The leaves of the trees are mainly compound with small leaf-
lets, but in many of them the leaflets are more than one square
centimeter in area. The number of trees with large simple
leaves is very small. The seasonal habits of foliage differ
widely in some of the common trees. A few of them are ever-
green (Cassia atomaria, Gochnatia arborescens, Sebastiania bilocu-
laris), a small number are winter deciduous (Prosopis glandulosa,
P. Palmeri), and a large number are drought deciduous (Jatropha
cinerea, Bursera laxiflora, B. microphylla, Cyrtocarpa edulis, Cer-
cidum sonorae). Many others are partly deciduous in the dry
spring months, the extent of defoliation doubtless depending on
the severity of the dryness at that season.

The most common trees and other tall plants in the Cape
forest below 1000 meters (3280 feet) are the following:

Lysiloma microphylla Esenbeckia flava
Jatropha cinerea Albizzia occidentalis
Cyrtocarpa edulis Gochnatia arborescens
Bursera laxiflora Haematoxylon brasiletto
Lysiloma candida Lemaireocereus Thurberi
Cercidium peninsulare Sebastiania bilocularis
Leucaena microcarpa Bursera microphylla
Cassia atomaria Pithecolobium tortum
Pachycereus pecten-aboriginum Plumeria acutifolia
Cercidium sonorae Bursera odorata
Prosopis Palmeri Yucca sp.

Small trees with soft wood and large shrubs from 2 to 5
meters in height are abundant and very important in contribut-
“ing to the physiognomy of the vegetation. The most prominent
species, including cacti, are the following:

Tecoma stans Lippia formosa
Viguiera tomentosa Viguiera deltoidea
Karwinskia Humboldtiana Opuntia cholla
Melochia tomentosa Turnera diffusa
Mimosa Brandegei Mimosa Xanti
Euphorbia Xanti Hypis tephrodes
Bourreria sonorae Alvordia fructicosa
Colubrina glabra Acacia flexicaulis
Ruellia californica Calliandra californica
Pithecolobium confine Fouquieria peninsularis
Gossypium Davidsonii Machaerocereus gummosus
Randia Thurberi Opuntia fuliginosa
Celosia floribunda Randia armata

Citharexylum flabellifolium

112 MADRONO [Vol. 4

The approximate distribution of the forest of the Cape region
is shown on the accompanying map (fig. 1). On the Pacific coast
it extends from sea-level to the borders of the mountain type of
forest, found above 1000 meters. A short distance north of
Todos Santos it gives way to the low desert scrub which charac-
terizes the outer edge of the Magdalena Plain. On the Gulf
Coast it covers all of the granitic outwash and certain favorably
located areas of volcanic outwash but is not found on the vol-
eanic hills near the coast south of La Paz. In the Sierra
Giganta it occupies the slopes of the mountain on the west and
the upper slopes on the arid eastern side. At the northern tip
of the Cape forest its occurrence at lower elevations is limited
to canyons and broad structural depressions in the lava fields.

The Magdalena Plain and the eastern coastal fringe south of
latitude 26° N. are desert in both vegetation and flora. South
of La Paz the character of the underlying rock and the derived
soils is closely correlated with the distribution of forest and
desert. Trustworthy inhabitants state that the east coast re-
ceives less rain than the interior as far north as Concepcion Bay,
a circumstance which adds to the aridity of the volcanic areas.
The Magdalena Plain is like the central region of the peninsula
in having little rain at any season. There is some morning fog
in April and May, and a narrow coastal strip is visited by strong
ocean wind varying in relative humidity from 55 to 65 per cent.
The region is distinctly unfavorable to the northward spread of
the Cape forest.

The number of plants which are common in the Cape Region
but absent from the desert of Baja California is large. The in-
filtration of the Cape vegetation by desert plants is considerable
as to the number of species but only locally important with re-
spect to their role in the vegetation. Larrea reaches the top of
the southern end of the escarpment but does not descend into the
plain of La Paz. Pachycereus, Lemaireocereus, Cercidium, Bursera
and other genera prominent in the desert are also frequent in
the Cape vegetation. Many of the common plants of the desert
are abundant on the volcanic areas in the Cape Region but only
sparingly represented in the Cape forest. On the volcanic areas
the height, spacing, types of plants and other physiognomic fea-
tures are identical with those of the desert areas far to the north,
and the composition of the vegetation is very similar.

Following is a list of the principal desert plants found in dry
and open habitats in the Cape forest. A few of these occur in
the desert only south of latitude 29° N. The remainder are
found nearly throughout the desert of Baja California and some
of them occur in southwestern Arizona also. The extent of the
northern range is indicated after each species.

Pachycereus Pringlei B.C. Brickellia Coulteri Ariz.
Bursera microphylla Ariz. Beloperone californica Ariz.
Lemaireocereus Thurberi Ariz. Opuntia cholla 29° N.

1937 | SHREVE: VEGETATION OF BAJA CALIFORNIA 113

Encelia farinosa Ariz. Machaerocereus gummosus B.C.
Fouquieria peninsularis 29° N. Calliandra californica B.C:
Solanum Hindsianum Ariz. Franseria magdalenae 29° N.
Trixis californica Ariz. Colubrina glabra 29° N.
Franseria ambrosioides Ariz. Condalia spathulata EXENZ,:
Bursera rhoifolia B.C. Simmondsia californica AUEIZ,
Pedilanthus

macrocarpus 29° N. Euphorbia tomentulosa B.C.
Jatropha cinerea B.C. Jatropha spathulata Ariz.

On the level plains of Sonora the desert merges gradually
into thorn-forest between latitude 27° and 28° N. In Baja Cali-
fornia there is uninterrupted desert as far south as latitude 26°
N.; it covers more than half of the peninsula thence south to lati-
tude 24° N., and small areas of it are found almost to the ex-
treme tip. The transition from desert to Cape forest is not a
matter of gradual change over many miles so much as the inter-
digitation of the two over rugged and varied country.

Differences of flora and of vegetation are not of the same
kind and are therefore difficult to compare. From a general
standpoint, nevertheless, it may be said that the Cape Region
and the adjacent mainland differ in vegetation fully as much as
in flora. The southern limit of desert is about three degrees
further south in Baja California than it is on the mainland. The
relationships of the flora of the Cape Region, as far as known,
are somewhat closer to Sonora than to Sinaloa. The flora of
the mainland, over a comparable area, is much richer than that
of the Cape Region, but the vegetation of the lowlands of the
Cape is very much more diversified than that of the lowlands of
Sinaloa and southern Sonora. In the latter region a single tree
is everywhere strongly dominant, and its commonest associates
are few as compared with the great variety of arborescent forms
in the Cape forest.

Desert Laboratory,

Carnegie Institution of Washington,
Tucson, Arizona, July, 1937.

BIBLIOGRAPHY

1. Bryant, W. E. The Cape Region of Baja California. Zoe, 2: 185-201.
1891.
Branpecer, T. S. A Collection of Plants from Baja California, 1889.
Proc. Cal. Acad. Sci. Ser. 2, 2: 117-232. 1889.
Flora of the Cape Region of Baja California. Proc. Cal.
Acad. Sci. Ser. 2, 3: 108-182. 1891.
. The Distribution of the Flora of the Cape Region of Baja
California. Zoe, 3: 223-231. 1892.
GRINNELL, JosePpH. A Distributional Summation of the Ornithology of
Lower California. Univ. Cal. Publ. Zool. 32: 1-300. 1928.
6. Netson, Epwarp W. Lower California and its Natural Resources. Mem.
Nat. Acad. Sci. 16. 1921.
7. Sureve, Forrest. Vegetation of the Northwestern Coast of Mexico. Bull.
Torr. Bot. Club, 61: 373-380. 1934.
8. ————_—_——.. Lowland Vegetation of Sinaloa. Bull. Torr. Bot. Club, in
press.

114 MADRONO — [Vol. 4

A NEW THALICTRUM FROM MOUNT RAINIER,
WASHINGTON

Harowtp St. JoHN

Thalictrum rainierense sp. nov. Perennial, producing a
slender offset at base; stem 30—48 cm. tall, glabrous, with 9-11
angles, the base short decumbent, clothed with brown marcescent
leaf sheaths; basal or sub-basal leaves usually 1 at anthesis, the
petiolar sheath 15-25 mm. long, membranous, brown, strongly

° 3mm

Fig. 1. a, stem and basal leaf; b, tip of stem and pistillate inflorescence;
c, fruit; d, achene.

ribbed, auricled at tip; petiole 4-5 cm. long, glabrous; blades
biternate, the petiolules 1-8 mm. long; leaflets 8-16 mm. long,
thin chartaceous, above dark green and glabrous, beneath pale
and glaucous and minutely capitate glandular puberulous, in out-
line suborbicular to broadly rhombic, the basal half entire, the
apex 1—2-times lobed with broad ovate or rounded lobes; cauline
leaves 38-4, the upper gradually reduced and shorter petioled;
staminate plants not seen; pistillate plants with the inflorescence
25-45 mm. long, loosely cymose, 3—5-flowered, glabrous; bracts
minute, suborbicular; pedicels 8-15 mm. long; calyx deciduous,
not seen; the 5-12 achenes stipitate at base the outer ones on a
stipe 0.2-1 mm. long, the inner on a stipe 1—1.8 mm. long, the

1937] JOHNSON: SEEDLINGS OF EUGENIA HOOKERI 115

body 5.5-6 mm. long, 1.2-1.7 mm. wide, obliquely slender fusi-
form, with about 9 prominent, longitudinal ribs; stigma 0.43
ime long. (Fig, 1.)

Perenne 30—48 cm. altum, caule glabro angulato, foliis biter-
natis, foliolis 8-16 mm. longis tenuiter chartaceis suborbiculari-
bus vel late rhomboideis subtus glanduloso-puberulis apicibus
lobatis, inflorescentibus foemineis cymosis 3—5-floriferis; acheniis
fusiformibus stipitatis.

Washington: in meadows, altitude 6000 feet, Mount Rainier,
August, 1895, C. V. Piper 2022 (type in State College of Washing-
ton Herbarium, Pullman). |

No similar species is known in the region. The closest rela-
tive seems to be TJ. stipitatum Rydb. (not T. stipitatum Rose,
1903), a native of the mountains of Colorado. It has the herbage
glabrous, and the achenes about 6 mm. long, and 2.5—3 mm. broad.
T. rainierense has the leaflets capitate glandular puberulous
beneath, and the achenes 5.5—6 mm. long, and 1.2—1.7 mm. wide.

University of Hawaii, Honolulu,
February 7, 1937.

SEEDLINGS FROM POLYEMBRYONIC SEEDS OF
EUGENIA HOOKERI

ArtHuR M. JoHNSON

In a previous paper’ the writer described an unusual poly-
embryonic condition found in the seeds of Eugenia Hookeri, a
common ornamental tree in the Los Angeles area. The question
naturally followed as to whether or not seedlings would develop
from these embryos and what form the seedlings would take.
The present paper deals with the results obtained from seeds
that were allowed to germinate in the soil under the parent tree.
The species produces an abundant crop of fruit annually in this
locality, and if the fallen fruits are allowed to remain on the
ground a good crop of seedlings will usually spring up.

The seedlings herein described were dug up on May 26, 1934.
The parent tree stands within a few feet of the north side of a
dwelling, where, except during the midsummer season, no direct
sunlight falls upon the ground beneath it. The soil is clayey
and is always moist, and frequently wet, especially on days
when the adjacent lawn is watered.

At the time these seedlings were collected numerous other
seedlings were growing in the ground beneath the parent tree.
Many more seedlings were examined than the ones here de-
scribed and figured. Seedlings have appeared annually in vary-
ing numbers since these were collected, though at the present

1 Johnson, A. M. Polyembryony in Hugenia Hookeri. Amer. Journ. Bot.
23: 83-88. 1936.

116 MADRONO [Vol. &

writing there are relatively few, owing perhaps to the recent
long cold spell.

One peculiarity of these seedlings is that they have never de-
veloped further than to a height of about a decimeter. The past
summer (1986) a number of seedlings, all less than a decimeter
in height, were transplanted to a more favorable location, in
drier soil, where they would be exposed to direct sunlight for
the greater part of the day. Up to the present they have made
no growth, although they are all as green and healthy looking as
when planted. It remains to be seen whether they will resume
growth after the advent of the warm season. |

The thirteen seedlings illustrated (Pl. XIX) were selected
from a representative lot of twenty-eight, which were growing
in scattered clusters in the shade beneath the parent tree. The
tallest specimen (Pl. XIX, fig. A) was 7.5 centimeters in height,
measured from the cotyledonary node to the uppermost visible
node. The most vigorous specimen (Pl. XIX, fig. C—drawn on
a larger scale for the sake of a clearer presentation of certain
details) measured 7 centimeters in height. Seedlings from the

EXPLANATION OF THE Figures. PLATE XIX

A. Normal seedling from a medium-sized embryo with unequal cotyledons.
Shoot 7.5 cm. in height.

B. Twin seedlings from a 3-cotyledonous embryo. Seedlings connate at
the cotyledonary node. Cotyledons very unequal; ¢, testa; a, b, c, cotyledons.

C. Vigorous seedling from the largest embryo of a seed. Shoot forking
at second epicotylary node. ‘Twin primary roots.

D. Seedling from a large embryo. Twin primary roots.

E. Interlocked seedlings (#, y) from two embryos of the same seed; x, two
unequal shoots from the epicotyl of the smaller embryo, the cotyledons of
which are a, b, c, and the primary root, r; y, well developed shoot from the
larger embryo, the cotyledons of which are d, e, f, and the primary root, s.

F. The smaller seedling of Fig. E, showing the two shoots, x, springing
from the epicotyl, the three unequal cotyledons, d, e, f (corresponding to d, e, f
in fig. E), and the forking primary root, r.

G. Opposite side of the seedling shown in fig. F, with the cotyledons cor-
respondingly labeled.

H. Part of the larger seedling, y, of fig. E, showing the three unequal
cotyledons (labeled correspondingly).

I. Normally developed seedling from one of the larger embryos of a seed,
with unequal polyhedral cotyledons.

J. Seedling from one of the smaller embryos, with twin primary roots.
Cotyledons unequal and polyhedral.

K. Seedling from a large embryo, with two unequal and arrested primary
roots.

L. Young normal seedling from a large embryo. Cotyledons nearly equal
in size but of irregular shape.

M. Part of a large seedling from a large embryo. Closely appressed
against its cotyledons is a small seedling from a minute embryo of the same
seed; a, epicotyl; 7, part of the primary root; c, the cotyledons of the larger
embryo, closely appressed against each other on one side of the seedling; s,
cotyledons of the smaller embryo.

1937]

JOHNSON: SEEDLINGS OF EUGENIA HOOKERI

Pirate XIX. SeEEepitines or Eucenta HooKkert.

117

118 MADRONO [Vol. 4

smallest embryos were but a few millimeters in height; one such
specimen, 12 millimeters in height, is shown (Pl. XIX, fig. J).

Of these twenty-eight seedlings, six had large cotyledons of
unequal size. In five others the cotyledons were small, and in
the remaining seventeen seedlings the cotyledons were of inter-
mediate size. The size of the cotyledons, however, did not ap-
pear to have had any bearing on the size of the seedlings. It
should be remarked in passing that the cotyledons are hypogeal,
and that the extreme differences in size and shape is due to their
close packing in the seed. Cotyledons of equal size in the same
embryo are not uncommon, although more frequent in smaller
embryos than in larger ones. Tricotyledonous embryos are also
not infrequent, and some seedlings from such embryos are herein
figured and described.

The primary root was in general well developed,—to the
length of the shoot or longer (Pl. XIX, figs. A, B, C). But in
some instances it was remarkably short, as if arrested early in
its development. In one small seedling in particular two such

primary roots were present, but were very unequal in length ©

(Pl. XIX, fig. K). There were several seedlings with twin pri-
mary roots of considerable length, more or less equally devel-
oped. (PI. XIX, figs. C, D, J.) | In these cases: only, a)isingie
shoot had developed from the epicotyl. In one instance, a
small tricotyledonous seedling interlocked with a larger one, twin
shoots arose from the epicotyl, and the primary root forked into
two approximately equal branches a short distance from the tip
of the hypocotyl. The three cotyledons and the twin shoots
suggest a condition of connate embryos (Pl. XIX, figs. E, F, G).
But a clearer case of twin seedlings, each with its distinct primary
root, was found, in which the twins were connate at the cotyle-
donary node only, and in this case there were also three cotyle-
dons present (Pl. XIX, fig. B). Interlocked seedlings, as might
be expected under the circumstances, were not uncommon, and
most of them were difficult to separate in order to determine
to what extent, if any, they were connate. It was especially dif-
ficult in the case of seedlings from the small embryos. In the
vigorous seedling already mentioned (Pl. XIX, fig. C) the shoot
forked at the second node above the cotyledonary node, the
branches being equally well developed.

Seedlings which showed none of the pecularities above de-
scribed were frequent. These may be said to be normal, except
that the cotyledons were frequently very unequal in shape and
size (Pl. XIX, fig. A, I, L). In every seedling examined the
lower leaves were minute scale-like organs.

University of California at Los Angeles,
March, 1937.

LPR

a

1937] WIGGINS AND STOCKWELL: FRANSERIA 119

THE MARITIME FRANSERIA OF THE PACIFIC COAST

Tra L. WicGins AND PALMER STOCKWELL

Since Nuttall described Franseria bipinnatifida in 1841 it has
been accepted generally as being distinct from F. Chamissonis
Less. In 1907 Dr. Harvey M. Hall suggested that F. bipinnatifida
was probably not specifically separable from F. Chamissonis.*
However, Dr. Hall and other botanists of the Pacific Coast con-
tinued to accord it specific rank, the segregation based primarily
on the twice or thrice pinnatifid leaves.

At the suggestion of Dr. L. R. Abrams of Stanford Univer-
sity experimental work was done to test the hypothesis that the
races mentioned are variants of one ecospecies.

Through the courtesy of the staff the plants were grown and
the cytological work was done at the laboratory of the Carnegie
Institution of Washington on the Stanford University campus.
Professor James McMurphy of Stanford University very kindly
furnished greenhouse space and care for some of the cultures.

Seventeen crosses were made. Five were F. Chamissonis on
F. bipinnatifida, five reciprocal, four F. bipinnatifida on F. bipinnati-
fida, and three F. Chamissonis on F. Chamissonis. In all cases
there was some variation in the F, progeny. When F. Chamissonis
and F. bipinnatifida were crossed the F, hybrids showed as much
variation in the small numbers raised as would be expected in F,
hybrids. One cross was carried on to the F, generation. The
male parent, Ff. Chamissonis, from Half Moon Bay was collected
by Wm. Hiesey; the female parent, F. bipinnatifida, was collected
by D. D. Keck (no. 2197) five miles north of Cambria. The
chromosome number of each was 2N= 36. Two hybrid offspring,
intermediate in leaf character, were chosen; one was used as the
male and the other as the female parent. The chromosome num-
ber of each was 2N=36 and N=18. Further, M, and M,, were
normal and regular. On October 5, 1935, 104 seeds were planted
in a flat from which 79 plants were harvested and pressed about
June 20, 1936. Of these, 11 had leaves entire, or nearly so; 36
had coarsely cut leaves; and 32 had finely cut leaves.

These tests, as well as field observations, indicate that in most
places the maritime Franseria is heterozygous and unstable. The
leaf form with associated characters is dependent on genetic con-
stitution, and this, of course, is dependent on chance. Maintain-
ing these forms as species does not seem to be justified as they
cross freely; one form swamping out the other at times, but both
being found from British Columbia to southern California. They
have the same chromosome number, both mitosis and meiosis are
normal, and cytologically they are indistinguishable. Therefore,
the following taxonomic treatment of these subspecies is pro-

posed.
1 Hall, Harvey M. Univ. Calif. Publ. Bot. 3: 121. 1907.

120 MADRONO [Vol. 4

1. Franseria CuHamissonis subsp. typica nom. nov. Franseria
Chamissonis Less. Linnaea 6: 507. 1831. Franseria Chamissonis
var. malvaefolia Less. Linnaea 6: 507. 1831. Franseria cuneifolia
Nutt.. Trans. Am. Phil. Soe. ser. 2, 7: 345. 1840. Fransena
Chamissonis var. cuneifolia Torr. & Gray, Fl. N. Am. 2: 298. 1842.
Gaertneria Chamissonis Kuntze, Rev. Gen. 339. 1891. Ambrosia
Chamissonis Greene, Man. Bay Reg. Bot. 188. 1894.

Leaves serrate to the cuneate base; bur channeled above.

Sandy coastal regions, San Clemente and San Miguel Islands,
and from middle California to Washington.

2. Franseria CuHamissonis subsp. bipinnatisecta comb. nov.
Franseria Chamissonis var. bipinnatisecta Less. Linnaea 6: 507.
1831. Franseria bipinnatifida Nutt. Trans. Am. Phil. Soc. II. 7:
344. 1840. Franseria Lessingii Meyen & Walp.; Walp. Nova
Acta Acad. Leop. Carol. 19: Suppl. 268. 18438. Gaertneria bipin-
natifida Kuntze, Rev. Gen. 339. 1891. Ambrosia bipinnatifida
Greene, Man. Bay Reg. Bot. 187. 1894. Franseria bipinnatifida
dubia Eastw. Proc. Calif. Acad. ser. 8,1: 117. 1898. Gaertneria
bipinnatifida dubia A. Heller, Muhl. 1: 6. 1900. Franseria bipin-
natifida villosa Kastw.; Rydb. N. Am. FI. 33: 26. 1922, as syno-
nym. Franseria villosa Rydb. N. Am, Fl. 33: 26. 1922.

Leaves once to thrice pinnatifid, bur ovoid but more slender
than in typica with spines sub-terete and usually only slightly
channeled above.

Range: British Columbia to Lower California. The South
American form seems to be this subspecies or a variant thereof,
and may be an introduction.

Stanford University, California.
February 10, 1937.

ON THE POLLEN OF THE MIMOSOIDEAE AND THE
IDENTITY OF THE SUPPOSED ALGA
PHYTOMORULA

Hersert F. Corpetanp

The state of California harbors a few native species of Mimo-
soideae, together with a large number of introduced species, in-
cluding about three of Albizzia and more than three-score of
Acacia. It may not be inappropriate to summarize what is known
about the remarkable clusters of pollen grains produced by some
of the members of the group. The existence of these clusters
is no new discovery: it is noted in several of the standard refer-
ence works (1, 8, 10, 11); it has been reported for Californian
material by Rowe (7); it has been known for more than a hun-
dred years.

I have not been able to confirm a reference to the writings
of Kohlreuter. The oldest paper which I have seen that de-
scribes these clusters is by von Mohl (4), who distinguished a

1937] COPELAND: POLLEN OF MIMOSOIDEAE 121

variety of types of clusters almost exactly as they are known
up to the present.

Rosanoff (6) added examples to the types already distin-
guished by von Mohl, and studied the development; his account
of the process in the majority of the species may be transcribed
into modern terms as follows. In each microsporanguim, all of
the archesporial cells except two become sterile. The two fer-
tile cells divide once, twice, or three times, forming clusters of
two, four, or eight microspore mother cells. Each of the latter
forms a tetrad of microspores. The outcome is the production,
by each anther, of eight clusters of pollen grains, each cluster
consisting of eight, sixteen, or thirty-two grains. (It is evident
that by homology each unit of the cluster is a pollen grain. We
should not regard the whole cluster as a single grain, though
some authors have done so.) As an anomalous exception, Rosa-
noff found in a few species clusters of twelve pollen grains; these,
it is evident, represent clusters of three pollen mother cells
derived from single fertile archesporial cells.

Wodehouse (11) brought to the description of these clusters
an extensive knowledge of the systems of grooves on the sur-
faces of pollen grains. He interprets the quadrate markings
found on the exposed surfaces of the grains of Acacia as repre-
senting his dodecacolpate (12-grooved) system. In the species
commonly known as Acacia Farnesiana, he finds the grains hexa-
colpate (6-grooved); this seems sufficient justification for his
excluding it from Acacia, under the old name Vachellia Farnesiana
Wight & Arnold.

What remains to be accomplished is a correlation of the
characters of the clusters and grains with the classification of the
group. Rosanoff remarked of the variety of types of clusters
that it “dient noch einmal als Bestatigung des alten Satzes von
Kohlreuter, dass die Aehnlichkeit der Pollenkérner nicht immer
mit dem Umgrenzung der Verwandtschaftskreise zusammen-
fallt.”” This in effect gave the taxonomists license to pay no
attention to the pollen grains; and they seem to have taken ad-
vantage of it. But Rosanoff wrote at a time when the limits of,
the most familiar genera, Mimosa, Acacia, and Albizzia, were not
understood; and I have encountered a number of facts which sug-
gest that when all of the genera are well understood the type
of pollen will be found uniform in each. Thus as noted above,
when Wodehouse found the Farnesian species to differ from
Acacia in a character of the pollen, he found also that it had long
before been excluded from Acacia on the basis of other charac-
ters. Again, Merrill removed the rain tree (a common orna-
mental in the tropics) from Pithecolobium, and placed it in an-
other genus as Samanea Saman; its pollen grains are in clusters
of thirty-two, instead of sixteen as in the only species of Pithe-
colobium I have examined. Most species of Inga seem uniform
in type of cluster; but von Mohl, dealing with I. anomala, and

122 MADRONO [Vol. 4

Rosanoff, dealing with J. tergemina, found in these species a quite
distinct type. I have myself found this type in Calliandra grandi-
flora; and I cannot but suspect that the peculiar species of Inga
might better be placed in Calliandra. But the re-shuffling of the
group cannot be attempted at this time. More than thirty genera
are recognized in the Mimosoideae; we know by observation in
California how numerous in species Acacia is; and a glance at
the literature shows that Inga, Pithecolobium, Enterolobium, and
Calliandra are comparably numerous.

The following outline of the pollen types is based primarily
on my own observations; the examples which I have seen are
marked with exclamation points. All the observations were
made on herbarium material. The pollen is most easily seen by
dissecting out and crushing the anthers of buds which are just
in the act of opening. From among the many examples offered
by the older authors I have cited but few, because I do not trust
the generic names they used.

I. THE GRAINS OR CLUSTERS NUMEROUS IN EACH ANTHER

Grains solitary: Leucaena glauca! Desmanthus spp. and Entada
spp., fide von Mohl, Rosanoff, and Wodehouse; all species
of Prosopis, fide Wodehouse; some species of Mimosa, fide
Rosanoff.

Grains in tetrads: Mimosa pudica and other species, also
Schrankia unciniata, fide Rosanoff.

Grains in octets: Schrankia sp., fide Rosanoff.

II. THE GRAINS IN CLUSTERS; CLUSTERS EIGHT IN EACH ANTHER

The tetrads tetrahedral, so that the grains lie in more than one

plane.

1. The grains interpretable as dodecacolpate, each having a
quadrate marking (sometimes seen as circular) on
the exposed surface.

a. Grains in octets: Acacia armata! A. tenuifolia (Pl. XX,
fig. 1)! and various other species, fide von Mohl and
Rosanoft.

b. Grains in clusters of twelve: Acacia rutaefolia, A. penta-
denia, and A. pulchella, fide Rosanoff.

ce. Grains in clusters of sixteen, the greatest dimension of
the cluster 80-50 microns. Throughout the family,
clusters of sixteen or more grains are always of a
lenticular form, half of the grains forming a circum-
ferential belt, while the rest are in two clusters, one
on each of the broad surfaces. The limits of the
tetrads are obvious, each consisting of two adjacent
circumferential cells and one cell from each of the
superficial groups. Acacia Baileyana (Pl. XX, fig.
2)! A. cultriformis! A. decurrens var. dealbata! A.
elata! A. longifolia! A. melanoaxylon! A. neriifolia! A.

1937] COPELAND: POLLEN OF MIMOSOIDEAE 123

podalyriaefolia! A. retinoides! A. verticillata! and vari-
ous other species, fide von Mohl and Rosanoff. The
only species outside of Acacia which is known to me
as probably belonging here is Archidendron Vaillantii;
Taubert’s figure (after von Mueller) appears to
show the quadrate surface markings.

2. The grains hexacolpate, the exposed surfaces divided into
three prominent protuberances; grains in clusters
of sixteen: Vachellia Farnesiana!

3. The grooves essentially obsolete, so that the grains appear
smooth.

a. Grains in clusters of sixteen, of which the greatest
dimension is 75-100 microns: Albizzia Acle! A. Juli-
brissin (Pl. XX, fig. 3)! A. Lebbek! A. lophantha! Inga
myriantha, fide Wodehouse; Pithecolobium dulce!

b. Grains in clusters of thirty-two, greatest dimension 90—
160 microns: Inga cordistipula, fide Taubert after
Bentham (actually seven tetrads, not eight, are
shown) ; I. edulis! I. spectabilis, fide Rosanoff (as an
individual variation, some clusters of thirty-six
grains) ; Samanea Saman! (Pl. XX, fig. 4).

Cells of the tetrad lying in one plane; tetrads always two, so that
the cluster is a flat plate of eight. The clusters are ex-
ceedingly large, to 250 microns long; the two ends are
not alike, so that the outline in surface view is lanceo-
late. Von Mohl reported a cluster of minute cells,
serving as a clinging organ, at the pointed end. Ro-
sanoff could see nothing but the naked point. My own
understanding of the situation is as follows. The
pointed end is, in the anther, pressed against a large
sterile cell. Under some circumstances it may break
completely free; under others, it may be found attached
to a scrap of cell wall. Inga anomala, fide von Mohl; J.
tergemina, fide Rosanoff ; Calliandra grandiflora! (Pl. XX,
fig. 5).

On the basis of these data, one may point out a few mistakes,

of observation or of interpretation, which have been published.

Rowe’s figure of pollen of Acacia is readily recognizable but
is not strictly accurate. It duly shows the quadrate protuber-
ances on the four cells occupying one surface of the cluster.
But as to the belt of circumferential cells, it shows protuberances,
not on the edges, but on the surfaces toward the observer. Von
Mueller’s figure for Archidendron Vaillantii shows the same fea-
ture, and is presumably in error in the same way.

Pope (5), referring to Acacia mollissima (the name is a
synonym of A. decurrens), describes “Grains . . . almost spheri-
eal; surface reticulated, reticulations forming large 4-sided
facets.” It is evident that she has mistaken the cluster of six-
teen grains for a single grain.

124 MADRONO [Vol. 4

Some twenty years ago, Kofoid (2) described and illustrated
certain sixteen-chambered structures found by him in a reservoir
in Berkeley during the month of March (when, according to
Rowe, Acacia pollen is most abundant). He looked about, natu-
rally, for something with a family resemblance to these struc-
tures, and there was nobody to call to his attention the Mimo-
soideae; he recognized Coelastrum as something similar, and de-
scribed his material as a new alga, Phytomorula regularis. More
recently, Smith (9) has reported and illustrated an additional
collection of Phytomorula (at Stanford University, according to a
personal communication from Dr. Smith). The two accounts
refer to things slightly different in size and shape, but within the
range of the size and shape of clusters of Acacia pollen, and con-
sisting of units arranged as in Acacia pollen. Kofoid’s illustra-
tion shows on each cell a minute dome-shaped protuberance;
Smith’s shows a large one; the protuberances as shown do not
agree well with each other, nor with the large quadrate protuber-
ances on the pollen grains of Acacia. I have not been able to ex-
plain these discrepancies; possibly they depend on conditions of
exposure. I am confident, however, that Phytomorula as pub-
lished respectively by Kofoid and Smith is the pollen of two
different species of Acacia.

Sacramento Junior College,
Sacramento, California, March, 1937.

LITERATURE CITED

1. Firrinc, H., and others. Eduard Strasburger’s Lehrbuch der Botanik
... 18 Auflage. Jena. 1931 (I have not studied the many editions
of this standard text, to determine in which one the clustered pollen
grains of the Mimosoideae were first mentioned. It was a later one
than the second (1895) but presumably earlier than the one here
cited).

2. Korom, C. A. Phytomorula regularis, a symmetrical protophyte related
to Coelastrum. Univ. Calif. Publ. Bot. 6: 35-40, t. 7. 1914.

3. von Mont, H. Uber den Bau und die Formen von Pollenkérner. Bern.
1834. non vidi.

4, ————————.. Sur la structure et les formes des graines de pollen [Ab-
stract, in French, of the above, by Leret]. Ann. Sci. Nat. 2e sér. 3:
148-180, 220-236, 304-346, tt. 9-11. 1835.

5. Port, M. A. Pollen morphology as an index to plant relationship. Part
II. Key to the families. Univ. Colorado Studies 15: 171-215. 1926.

6. Rosanorr, S. Zur Kenntniss des Baues und der Entwicklungsgeschichte
des Pollens der Mimoseae. Jahrb. wiss. Bot. 4: 441-450, tt. 31, 32.
1865.

7. Rowr, A. H. A study of the atmospheric pollen and botanic flora of the
east shore of San Francisco Bay. Journ. Lab. and Clin. Med. 13:
416-439. 1928.

8. ScunarF, K. Vergelichende Embryologie der Angiospermen. Berlin. 1931.

9. Smiru, G. M. Fresh-water algae of the United States. New York. 1933.

10. Tausert, P. Leguminosae. In A. Engler and K. Prantl. Die Natiirlichen
Pflanzenfamilien 3°: 70-385. 1894.

11. Wopenovuse, R. P. Pollen grains, their structure, identification and sig-
nificance in science and medicine. New York. 1935.

1937] COPELAND: POLLEN OF MIMOSOIDEAE 125

Piate XX. CLUSTERS OF POLLEN GRAINS OF CERTAIN Mimosomnear. Fig. 1.
Acacia tenuifolia x 800. Fig. 2. Acacia Baileyana xX 800. Fig. 3. <Albizzia Juli-
brissin X 800. Fig. 4. Samanea SamanX 400. Fig. 5. Calliandra grandiflora
x 400.

126 MADRONO | [Vol. 4

NATHANIEL LYON GARDNER
(1864-1937)

There has been removed from our society Nathaniel Lyon
Gardner, Professor of Botany, Emeritus, of the University of
California. Born in Keokuk, Iowa, on February 26, 1864, he
passed away at his home in Berkeley, California, on Sunday,
August 15, 1937. His death resulted from a major cerebral
hemorrhage after several minor attacks, especially in the optic
tract, through a number of years, and which had disabled him
and prevented him from carrying on his studies for several
months. He was married in 1915 and is survived by his wife,
Edith Jordan Gardner, daughter of the late David Starr Jordan,
eminent educator and ichthyologist.

The early education of Dr. Gardner was carried on in Iowa,
but, after he began to teach school, he removed to Island County,
Washington, and continuing his interest in the flora which he had
begun in Iowa, sent specimens of various groups of plants to
eminent authorities for determination. It was in April, 1897,
that while teaching school on Whidbey Island, Washington, he
wrote to the writer, asking him as to whether he would be willing
to name some specimens of the marine algae of the shores of that
island. The request came at a crucial time in the writer’s work,
when the questions of the algae described from the Puget Sound
region and north were becoming critical in his own studies.
Nothing had been done in the way of collection or study in this
region for about thirty years. It is evident that an enthusiastic
and appreciative answer was dispatched and there began a scien-
tific cooperation which has lasted during these past forty years.
Through this association much has been done towards making
known the rich and peculiar marine flora of the Pacific coasts of
North America and work was still actively going on until the
afflictions of eye and general bodily health put an end to our
mutual effort. In fact, Gardner was carrying on the main work
on Pacific Coast algae while the writer pursued more and more
studies on the tropical marine floras of the Indo-Pacific region.

In his attempt to prepare himself for better research work,
Gardner, soon after the initiation of our correspondence, took up
residence at the University of Washington, from which institu-
tion he received the degree of bachelor of science in 1900. From
1900 to 1906 he acted as Assistant in Botany at the University of
California, advancing to the degree of master of science in 1903
and to that of doctor of philosophy in 1906. Leaving the Uni-

versity of California in the latter year, he became Head of the |

Department of Biology in the Polytechnic High School of Los
Angeles. He returned to the University of California in 1909-
1910 as Acting Assistant Professor of Botany, and became regu-
lar Assistant Professor in 1913. In 1923 he was appointed Asso-

Maprono VotumE IV, Pirate XXI

Pirate XXII. Naruaniet Lyon GARDNER.

1937] SETCHELL: NATHANIEL LYON GARDNER 127

ciate Professor of Botany, becoming Emeritus in 1934. From
1920 to 1984 he served as Curator of the Herbarium of the Uni-
versity of California; in 1934 he became Curator, Emeritus.

The scientific activities of Dr. Gardner were largely devoted
to the algae and he was primarily a phycologist, first, and always
exceedingly active in the field, collecting both the marine species
and those of the fresh waters. His specimens were selected and
prepared with great care and discrimination, as well as in large
numbers for each collection. The fresh-water species, particu-
larly those of the blue-green algae, were grown in his laboratory,
especial pains being taken to obtain cultures purely or predomi-
natingly of one species and under normal conditions. Many of
his cultures of blue-green algae were carried on for years and his
experience led him to reject the ideas of even moderate poly-
morphism among members of this group, an idea so readily
assumed by many writers.

From his abundant collections and his careful study of them,
his conceptions of species were narrow rather than broad and he
could demonstrate from his ample material his precise idea of
amplitude of variation and more particularly of its limits. His
very last studies were concerned with the Pacific North American
species of the genus of red algae, Iridophycus, and his extensive
field studies were correlated with no less ample studies of their
histology and development.

He was a world authority on the blue-green algae and pub-
lished a series of cytologic and taxonomic studies such as his
“Cytological Studies in Cyanophyceae” (1906) and his “New
Myxophyceae from Porto Rico” (1927).

His studies on various green algae, particularly marine spe-
cies, were numerous and thorough and he added much to both
our general knowledge and to our more accurate knowledge of
the green algae of our Pacific coasts.

Of the brown algae, he made extensive studies of the species
of our coasts, detecting, describing, and figuring an incredible
number of minute epiphytic forms, all hitherto undescribed. He
also studied the large forms of the brown algae in a similar
fashion, particularly the Fucales (or rockweeds), collecting abun-
dantly of them from Sitka, Alaska, to Ensefiada, Mexico. His
“Variations in nuclear extrusion among the Fucaceae” (1910)
and his “The genus Fucus” (1922) are only two of his outstand-
ing contributions. He also described two new fossil Fucales from
the Miocene of California.

Among the red algae, he felt less at home, but nevertheless
he made great progress and was becoming more confident in
general, while in particular genera he was perfectly equipped.

His work on fungi, while considerable, was only incidental.
_He published only one paper dealing with them but he was a
most successful collector of hypogaeous forms, such as the Tuber-
ales, Hymenogastrales, etc., and contributed much that was novel
in his collections.

128 MADRONO [Vol. 4

From 1903, much of his publication was done conjointly with
the writer but the greater part of the detailed work was done by
him. There are, however, approximately thirty larger and more
important papers for which he was willing to assume the sole
responsibility.

At his death he was working hard to complete the final vol-
ume of “The Marine Algae of the Pacific Coast of North Amer-
ica,’ a work including the practical monographing of a number
of large polymorphous genera of difficult red algae of wide dis-

tribution. His thoroughness and patience showed at their best in .

this work.

As a teacher, he was quiet and meticulous, not so successful
with large classes but, with small groups, his pupils received the
utmost attention and serious students appreciated his methods.

He was assiduous in exploring and devising methods for the
culture of his algae, for their proper preservation for future
examination, and for all methods of microtechnique, fixing, stain-
ing, sectioning, as well as obtaining the best possible optical
demonstration of their habit and structure.

Personally, Dr. Gardner was of a retiring disposition, content
to carry on his work and his studies without “fuss or flurry.” He
was always willing to interrupt his own work to help a worthy

beginner or more advanced student and gave freely of his own.

resources, both material and intellectual. His was not an enthusi-
astic nature, but rather deliberate, weighing the possible adverse
features; but his friendship, once given, was strong and enduring,
not of outward demonstration, but of faithful devotion.—W. A.
SETCHELL.

A NEW PENSTEMON FROM THE CHARLESTON
MOUNTAINS, NEVADA

Ira W. CLokey

Penstemon Keckii sp. nov. Herba perennis maximam partem
glaucescens; rhizoma ramosa; caules erectiusculi diluto-virides
vel interdum purpureo-tincti, infra glabri vel minute scabridi,
supra usque ad inflorescentem minute sparseque stipitato-glandu-
losi; folia pallido-viridia integra glabra, margine plus minusve
scabrido; folia basalia atque ea surculorum sterilium 2-6 cm.
longa oblanceolata, apice obtusa vel rotundata, in petiolum
alatum gradatim attenuata; folia caulina lanceolata sessilia;
racemus angustus; pedicelli 3-4 mm. longi sparse glanduloso-
puberulenti radiati demum sursum curvati; sepala 4-6 mm. longa
sparse glanduloso-puberulenta ovata acuta vel obtusa, margine
searioso; corolla 17-23 mm. longa, faucibus 6 mm. latis, infundi-
buliformis aequibilabiata, extus glandulosa, intus basi labri su-
periori excepto, eglandulosa, limbo atrocaeruleo, labro inferiore
ad medium inciso, pilis albis complanatis faucibus prominente

1937] CLOKEY: NEW PENSTEMON FROM NEVADA 129

barbatis; stamina didynama, staminibus superioribus corollae
faucis excedentibus, inferioribus inclusis; loculis antherarum 1.5
mm. longis late divaricatis fulvis glabris, de apice ad basin dehis-
centibus non confluentibus, suturis denticulatis; staminodium
staminibus inferioribus paullo brevius, apice dilatum pilis luteis
rigidiusculus 0.6 mm. longis dense barbatum; capsula ovata acuta
10-12 mm. longa.

Perennial with branching rootstock; herbage usually glauces-
cent; stems commonly 8-15 em. high but up to 25 cm., not strictly
erect, light green or sometimes purplish tinged, lower part gla-
brous or minutely scabridulous, becoming minutely and sparingly
stipitate-glandular towards and within the inflorescence; leaves
light green, entire, glabrous excepting the more or less scabridu-
lous margin; basal leaves and those of the sterile shoots 2-6 cm.
long, oblanceolate, obtuse or rounded at the apex, very gradually
reduced to a winged petiole; cauline leaves in two or three pairs,
sessile, lanceolate; inflorescence a narrow raceme occupying 14 to
22 of stem; pedicels sparingly glandular-puberulent, radiating
then curving upwards so that the flowers are erect, 3-4 mm. long;
sepals sparingly glandular-puberulent, ovate, acute or obtuse,
scarious margined, 4-6 mm. long; corolla deep blue 17-23 mm.
long, 6-9 mm. wide at throat, funnel-shaped, the tube exceeding
the calyx, glandular externally, glandular within only at base of
upper lip; corolla-limb of two equal lips, the lower prominently
bearded at junction with throat with flattened white hairs, the
lobes cleft about 14 the length of the lips; upper pair of stamens
exceeding throat, shorter pair included; anther sacs widely divari-
cate, buff, 1.5 mm. long, glabrous, dehiscent from the distal apex
essentially throughout, not confluent, the suture denticulate;
sterile filament reaching orifice, dilated apically, densely bearded
dorsally for most of its length with relatively short (0.6 mm.
long) stifish yellow hairs; capsule ovate, acute, 10-12 mm. long.

Deep gravelly, brushy meadow in the bed of Lee Canyon,
. Charleston Mountains, Clark County, Nevada, altitude 2570

meters, July 3, 1936, Clokey 7312 (type, Clokey Herbarium, South
Pasadena, California; cotypes being distributed in Exsiccatae
Grayanae). Other collections from the Charleston Mountains
which represent Penstemon Keckit are: Lee Canyon, August 1,
1935, Clokey 5579 (topotype), Charleston Peak, altitude 3150
meters, August 8, 1935, Clokey 5592, Rainbow Falls, altitude 2670
meters, July 27, 1936, Clokey 7311.

This is essentially a subalpine species and is widely but spar-
ingly scattered either on open hillsides or associated with Pinus
aristata Engelm. It extends, however, some distance below the
subalpine zone. At the Lee Canyon station, where it is fairly
abundant in an area about 200 yards across, the plants average
somewhat larger than those in the subalpine zone; here Penstemon
Keckii is associated with Pinus scopulorum (Engelm.) Lemmon.

130 MADRONO [Vol. 4

Penstemon Keckii belongs to the subgenus Glabri. Dr. Keck
suggests that this plant “is most closely related to P. speciosus
Dougl. which, in the broad sense, extends from arid eastern
Washington, southward through Oregon and the Sierra Nevada
to Mount Pifios in California and southern Esmeralda County in
Nevada.” In addition to being widely separated geographically
from P. speciosus, P. Keckii is smaller and occurs at a higher eleva-
tion. The following key may serve to separate the two species:

Inflorescence glandular, corolla glandular without and

within, copious beard at base of lower lip, sterile

Stamen: ‘strongly bearded yo ii. 50.2 2a tee ere P. Keckii
Inflorescence and corolla not glandular, the latter some-

times with a sparingly bearded throat and sterile

Stamen” bis. 3). Peas cua eee en erate nee eee P. speciosus

In appreciation of the work he is doing with western Pen-
stemon, it is a pleasure to name this species for Dr. David D.
Keck of the Carnegie Institution of Washington at Stanford
University.

South Pasadena, California,
June, 1937.

A NEW CALIFORNIAN SPECIES OF BRODIAEA

Rosert F. Hoover

Brodiaea appendiculata sp. nov. Cormo magno, sine pro-
pagulis, cum tunica crassa fibrosa circum scapum foliaque ex-
tendente; foliis 2—4, angustis; scapo 10-40 cm. alto; perianthio
infundibuliforme; tubo perianthii cylindrico, basi rotundo, 8-10
mm. longo, in fructo membranaceo; segmentis rectis, 15-20 mm.
longis, exterioribus oblongis, acutis, interioribus latioribus ob-
tusis; staminodiis linearibus, 8-12 mm. longis, obtusis vel retusis,
margine involutis undulatis; filamentis 4-5 mm. longis, margine
alatis, apice biappendiculatis; appendiculis linearibus, 3-5 mm.
longis, undulatis; antheris 7-8 mm. longis; capsula subglobosa,
apice acuta; seminibus in quoque loculo circa 5.

Corm large, deep-seated, without offsets, with a heavy coat of
brown fibers which extends as a sheath around the subterranean
portion of the scape and leaves; leaves 2—4, narrow; scapes stout,
1-3 from a corm, 10—40 em. tall; umbel 3-10 flowered, the pedi-
cels 3-9 cm. long, widely divaricate in age; perianth funnelform;
perianth tube cylindric, rounded at base, green, 8-10 mm. long,
in fruit membranaceous, finally brittle; segments straight (that is,
not recurved), 15-20 mm. long, purple with dark mid-vein, the
outer oblong, acute, the inner broader and obtuse; staminodia
purple with white tips, linear, obtuse or retuse, 8-12 mm. long,
with involute and undulate margin, approximate around the
anthers; stamens shorter than staminodia; filaments 4-5 mm.

1937] HOOVER: NEW CALIFORNIAN SPECIES OF BRODIAEA 131

long, purple, wing-margined, bearing at apex two appendages
which extend along the backs of the anthers; appendages linear,
undulate, pale purple or white, 3-5 mm. long, about half as long
as the anthers; anthers 7-8 mm. long, deeply notched at both
ends; capsule subglobose with acute apex; seeds about five in
each cavity.
Type: Warnerville, Stanislaus County, April 29, 1936, Hoover
1040.

a B i d

Fig. 1. Brodiaea appendiculata Hoover. Outline drawings based on fresh
plants: a, external view of perianth x1; b, dorsal view of staminodium x 2;
c, side view of staminodium X 2; d, dorsal view of stamen <3; e, mature cap-
sule X 1.

Brodiaea appendiculata is common in the lower Sierra foothills
and on the bordering valley plains from Sacramento County. to
Fresno County. It occurs also, but is apparently rare, in the
central Coast Ranges.

Herbaria indicated in the citations below are abbreviated as
follows: University of California (UC); California Academy of
Sciences (CA); herbarium of W. L. Jepson (J); Dudley Her-
barium of Stanford University (S).

Specimens examined. Sacramento County: Sacramento, May,
1919, Georgia Bentley (S), Hoover 1130 (J). Amador County:
Jepson 9957, 9965 (J). Calaveras County: Burson, Jepson 9954
(J); Jenny Lind, May 14, 1923, W. P. Steinbeck (CA). San
Joaquin County: Linden, May, 1896, Gunnison (UC); Peters, E.
E. Stanford 952 (S); Farmington, Hoover 1058 (UC). Tuolumne
County: Sonora, April 10, 1925, E. A. Green (S); open wooded
hill slopes near Bear Creek, Mrs. W. J. Williamson 49 (CA, S).
Stanislaus County: Warnerville, April 29, 1936, Hoover 1040
(type J, isotype UC); Oakdale, Hoover 1020 (UC); Montpellier,
Hoover 590 (UC). Merced County: Snelling, Hoover 964 (UC) ;
Merced Falls, Eastwood 4884 (CA); Merced, Hoover 1079 (UC);
dry rocky soil, edge of foothills, San Joaquin Valley, J. T. Howell
9 in part (CA). Madera County: Madera, Hoover 979 (UC).
Napa County: Napa Valley, Jepson 9981 (J). San Mateo County:
Cedro field, Stanford University, May 31, 1922, Bacigalupi (S).

132 MADRONO [Vol. 4

This species of the subgenus Hookera has long been known to
the author, but since the characteristic stamen appendages were
overlooked, it was thought to be a form of Brodiaea synandra
(Heller) Jepson. As a distinct species it was first recognized in
the field by W. L. Jepson, who made four collections of it in 1923
and at that time noted its distinctive characters. (W. L. Jepson,
Field, Book, vol..39: pp. 171, 178571775" 191,, 199), acs) eee
presence of appendages on the stamens has been noted previ-
ously only in B. stellaris Wats., a species occurring locally in the
North Coast Ranges, which has broad appendages very differ-
ent from the slender undulate ones of B. appendiculata. The short
capsules serve to distinguish B. appendiculata from both B. syn-
andra and B. stellaris, as well as from other species with which
it might be confused. The corms are more deeply seated and
have heavier sheathing coats than in other species of Brodiaea.
The horizontally spreading fruiting pedicels also, are distinctive;
among other species of the genus they have been observed only in

B. minor (Benth.) Wats.
University of California, Berkeley,
May 15, 1936.

REVIEW

Flora of southeastern Washington and of adjacent Idaho. By Haro.p
St. Joun. Students’ Book Corporation, Pullman, Wash-
ington. 1987. 581pp. 14figs. Imap. Cloth, $3.50;
paper, $3.00.

While Thomas Howell was bringing to conclusion his pioneer
Flora of Northwest America, in 1901, there appeared a modest
volume by Charles V. Piper and R. Kent Beattie, Flora of the
Palouse Region, treating the 668 species and varieties found within
a radius of 85 kilometers from Pullman, Washington. The
authors expanded this, in 1914, into A Flora of Southeastern Wash-
ington and Adjacent Idaho, with descriptions of 1,141 species and
varieties occurring naturally in the easternmost counties of Wash-
ington, from Spokane to Walla Walla, and a narrow strip of
neighboring Idaho.

The present work is a lineal descendent of these earlier floras
and frankly based upon them, covering the same area as the 1914
work, but it is not, in any sense, a compilation. Every descrip-
tion is original, drawn from the writer’s wide field experience,
or from personal consultation of specimens in numerous Ameri-
can and European herbaria. The preliminary study and writing
has occupied seventeen years and the book affords ample evi-
dence of the pains taken to make it clear, complete and in line
with the most recent monographic treatments. <A total of 1,473
species and subdivisions of species receives recognition and
description.

1937] REVIEW 133

St. John’s flora embraces most of the desirable features to be
found in a composite picture of the more recent manuals from the
frontispiece map showing life zones in color, through the unusu-
ally complete glossary to the useful “Explanation of Authors’
Names.” The format follows the popular style inaugurated in
W.L. Jepson’s Manual of the Flowering Plants of California, but the
genera and species are conveniently placed in strictly alphabeti-
cal order. A very clear and concise account of the physiography,
the climate and the vegetational features of the area precedes the
taxonomic treatment, but one might wish for more information as
to the component floristic elements, their affinities and probable
origin. C. Hart Merriam’s “life zones” are accepted as being the
best available indices of major plant environments.

The keys will, I think, evoke chief admiration! There are
keys from one end of the book to the other and they approach
mechanical perfection. The key to the families is helpfully illus-
trated with cuts of structures which usually confuse beginning
students. Salix and Antennaria boast keys to the staminate plants
and keys to the pistillate; the family Umbelliferae and the genus
Cogswellia are provided with separate keys to the plants in flower-
ing and in fruiting condition.

Charles Piper Smith is joint author of the treatment of
Lupinus and J. H. Barnhart assisted with the bibliography of
authors. Otherwise, the book is entirely its author’s own and
reflects his ideas and judgments throughout. It will be noted
that more of Wilhelm Suksdorf’s work has been taken cognizance
of—even to the extent of accepting five of his Amsinckia species—
than has been usual with American botanists. One recalls that
St. John not only knew Suksdorf personally but that he was the
man responsible for obtaining his priceless collection for the
State College of Washington.

The original plan was to call the flora “Edition II” of Piper
and Beattie’s later book and the 1914 work is freely cited as
“Edition I.” However, St. John subsequently decided to omit
the earlier authors’ names and altered their title slightly, but
continued to refer, rather ambiguously, to “Ed. I.’ Another
title would have been preferable for the present work, and the
idea was considered and discarded, although the use of such a
name as “manual” might have made the similarity in names less
confusing.

A tabulation of names reveals the following statistics: new
species, 17; plants “doubtless new” but not described, 6; new
varieties, 7; new forms, 7; new combinations, 39; new names, 4.
The author states, by way of preface, ““The writer does not sub-
scribe to the recognition of minute genera and species . . . those
minor elements so frequently announced as species by recent
American botanists. The writer's concept .. . is not materially
different from that of Piper.’ However, Piper’s specific concept

134 MADRONO [Vol. 4

was vastly different when he proposed 48 new species of Allo-
carya from that when he wrote The Flora of Washington; St.
John’s treatments show a comparable unevenness. In his treat-
ment of Rosa, St. John recognizes 8 species, 3 varieties and 3
“doubtless undescribed” species, and of these 4 species and 1
variety are described as new. George Neville Jones, during a
recent study (Madrofio 3: 120-135, 1935) saw the type collec-
tions of all of these but accepted only 9 species and 6 varieties of
Rosa for the entire state of Washington.

The inclusion of more than forty formae, nearly half of them
color-forms, is a precedent that few western botanists are likely
to follow. That these may have a place in monographs seems
defensible, but they seem completely out of place and pedagogi-
cally undesirable in a manual. Probably no one will criticize the
adoption of the Englerian arrangement but not all will agree
with St. John that the Engler and Prantl system “‘seems the best”
arrangement of plant families.

However, the points taken issue with here are largely matters
of individual preference and reflect the influence of one taxonomic
school. St. John has been highly successful in producing a thor-
oughly workmanlike, up-to-date and usable flora for southeastern
Washington, which will be of value to botanists everywhere but
will be an especial boon to those teaching systematic botany
within the area covered.—LIncoLn CoNnsTANCE.

NOTES AND NEWS

Word has been received from Japan that Dr. William A.
Setchell of the Department of Botany, University of California,
Berkeley, was elected an honorary member of the Botanical Soci-
ety of Japan at the general meeting held in Sapporo on July 29,
1937.

Dr. Francis W. Pennell, curator of plants at the Academy of
Natural Sciences, Philadelphia, spent the months of June, July,
and August collecting Scrophulariaceae principally in Montana,
Idaho, Oregon, and Washington. An especial effort was made to
follow the trail of Lewis and Clark through Idaho, and so to
determine the type locality of Penstemon fruticosus (Pursh)
Greene.

Miss Sarah C. Dyal, who is studying the Valerianaceae at
Cornell University, has been visiting Pacific Coast institutions
this summer. She assisted in the University of Wyoming summer
school, and plans to spend the winter at Oregon State College, at
Corvallis. In the spring, Miss Dyal hopes to gain further field
knowledge of the genus Plectritis in California and the Northwest.

The Associate Students’ store, University of California, Berke-
ley, reports that the Manual of the Flowering Plants of California,

y

1937] NOTES AND NEWS 135

by W. L. Jepson, is now out of print. It is to be hoped that a
new and revised edition of this valuable work will be soon forth-
coming.

Dr. Artemio V. Manza, who received the degree of doctor of
philosophy from the University of California in May, 1937, left
Berkeley on August 11, 1987, for Canton, China, where he has
accepted a position as Assistant Professor of Botany in Lingnan
University. Dr. Manza came to this country about ten years ago
from the Philippine Islands, first studying plant pathology and
then specializing in phycology. He has done considerable work
on the jointed corallines, a large group of red algae, recently
publishing a revision of the genera in this group.

On September 8, 1937, Mr. and Mrs. Carl W. Sharsmith left
Berkeley for Pullman, Washington, where Mr. Sharsmith will
take up his duties as instructor in the Department of Botany and
Curator of the Herbarium, State College of Washington. For
the past two years he has been teaching assistant in systematic
botany at the University of California, Berkeley, and for the past
year Mrs. Sharsmith has been instructor in biology at Mills Col-
lege, California.

Mr. Joseph A. Ewan has accepted a position in the Depart-
ment of Biology, University of Colorado, Boulder. For several
years he has been at the University of California, Berkeley,
where he assisted Dr. W. L. Jepson with the Flora of California.

Mr. F. Raymond Fosberg, recently on the faculty of the Uni-
versity of Hawaii, has received a fellowship at the Morris Arbo-
retum, University of Pennsylvania. Before leaving for the east
in August, 1937, Mr. and Mrs. Fosberg spent several days visit-
ing herbaria in Los Angeles and in the San Francisco Bay region.

A survey of the trees of Amador County has recently been
completed by Mr. Guy Towsley, principal of schools in Jackson,
California. The work was carried on under the guidance of Dr.
E. E. Stanford, College of the Pacific, Stockton, California, and
is filed in the library of that institution.

Dr. Ruth Harmon has been appointed instructor in bacteri-
ology and botany at Mills College, California. Dr. Harmon, who
received the degree of doctor of medicine at Friborg, Germany,
will also assist in the Mills College clinic.

Dr. Walter Robyns, Director of the State Botanical Garden,
Brussels, Belgium, visited the University of California on Sep-
tember 10, 1937. He is making a study of American and Euro-
pean herbaria and botanic gardens with a view to the reestablish-
ment of the garden and herbarium at Brussels as the growth of
that city has made it necessary to find a new location for the

136 MADRONO [Vol. 4

garden. Dr. Robyns has been commissioned by the Belgian gov-
ernment to visit our national parks in the interest of further de-
velopment of the Pare National Albert at Kivu, Belgian Congo.
On his trip west, he visited the parks in the northern part of the
United States. On September 21 he sailed for the Hawaiian
Islands where he will spend about ten days. Upon his return to
this country, he will complete his tour of the national parks.

The following recently published monographic treatments are
of interest to botanists of western North America:

“The genus Youngia,’ by Ernest B. Babcock and G. Ledyard
Stebbins, Jr. Carnegie Institution of Washington, Publication
No. 464, 106 pages, 5 plates, 31 text figures, August 19, 19387.
This group of Asiatic composites closely related to Crepis has
been found on cytological and taxonomic evidence to be distinct
from that genus. Twenty-seven species, grouped under six sec-
tions are recognized. In the introduction the authors discuss the
following subjects: taxonomic history of the genus, criteria of
classification, relationships of the genus and of the sections,
geographic distribution.

“The North American species of Rumex,” by K. H. Rechinger,
Jr. Field Museum of Natural History, Publication No. 386, 151
pages, 25 plates, June 24, 1937. Forty-seven species are recog-
nized: these are treated under three subgenera; Acetosella, 2
species; Acetosa, 4 species; Lapathum, 41 species. Under this
latter group two new sections are proposed: Axillares, 18 species;
Simplices, 23 species. In the introduction Dr. Rechinger dis-
cusses the distinctive characters and geographic distribution of
section Axillares which has its principal area of distribution in
North America. Users of the volume will appreciate the con-
sistently clear statements as to how the author’s interpretations
of certain specific units differ from those of current floras and
manuals.

“The genus Bidens,’ by Earl Edward Sherff. Field Museum
of Natural History, Publication No. 388, 346 pages, 88 plates,
August 31,1937. The introduction comprises a historical survey
of the taxonomy of the genus and mention of work done with
reference to it in such fields as morphology, histology, ecology
and relation to insect pollination. The systematic treatment
includes: description of the genus; synopses of the fourteen rec-
ognized sections; descriptions of species; citations of specimens
and comment. All keys, descriptions and synopses are in Latin.
In this cosmopolitan group of composites two hundred thirty-
three species are recognized and keyed out; of these, one hun-
dred one are treated systematically in Part I. The major sec-
tions of the key are geographic, as follows: Pacific islands; North
and Central America and the West Indies; South America; east-
ern hemisphere exclusive of Africa; Africa, an arrangement which
adds greatly to the usefulness of the monograph in any particular
region.

Complete Your Files!

MADRONO

A West American Journal of Botany

“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges.

Volume I, 1916-1929. . . $5.00
Volume II, 1930-1934 .. 5.00
Volume III, 1935-1936 . 5.00
Single numbers....... 0.75

There is only a limited supply of Vol-
umes I and II.

Address all communications
and orders to:

Dr. David D. Keck

Carnegie Institution of Washington
Stanford University, California

Pr gangrene “ae
ne tein BAM FB ier

f % 7 . J
‘eae Poe FP Rae tae
(as PUN OR EE FPR aD Pf TFs Ulm mm,

WOLUME IV = ,“y “OS. NUMBER 5

MADRONO

A WEST AMERICAN JOURNAL OF
BOTANY

we

Contents

wo New Species or LINANTHUS FROM WesTERN NortH America, Herbert
REMEMEROIB VAN SP Me SKU Aa ee TMb on ie MRK SON Sig SUR NARS i RGAE Se 157

VOTES ON Some San Dieco County Enpemics, Frank F. Gander .......... 163

[ae Catirorn1a Nutmec TREE in Curtivation, Albert Wilson ........... 166

Warren, The Plants of Mt. Rainier National Park, Washington (Lincoln
TUS eV) NBN Ga PREP ec BR RRA Ae eae OR 167

Published at Lime and Green Streets, Lancaster,
Pennsylvania

January, 1938

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asrams, Stanford University, California.

Dr. Lincotn Constance, University of California, Berkeley.

Dr. H. F. Copetanp, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Business Manager—Dr. Davm D. Kreck
Lime and Green Sts., Lancaster, Pennsylvania
or
Carnegie Institution of Washington
Stanford University, California

Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

Established 1916. Published quarterly. Subscription Price $2.50 per year.
Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17,
complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num-
bers $0.75.

Papers up to 15 or 20 pages are acceptable. Longer contributions may
be accepted if the excess costs of printing and illustration are borne by the
contributor. Articles may be submitted to any member of the editorial board. |
Manuscripts may be included in the forthcoming issue provided that the
contributor pay the cost of the pages added to the issue to accommodate his —
article. Reprints of any article are furnished at a cost of 4 pages, 50 copies
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at $1.50 per hundred. Reprints should be ordered when page proofs are
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Published at Lime and Green Streets, Lancaster,
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CALIFORNIA BOTANICAL SOCIETY
President: Dr. F. W. Foxworthy, Berkeley, California. First Vice-Presi-

dent: Dr. Ira L. Wiggins, Department of Botany, Stanford University. Sec-

ond Vice-President: Dr. Philip A. Munz, Department of Botany, Pomona
College, Claremont, California. Treasurer: Dr. David D. Keck, Carnegie In-
stitution of Washington, Stanford University. Secretary: Miss Ethel Crum,
4004 Life Sciences Building, University of California, Berkeley.

Annual membership dues of the California Botanincal Society are $2.50,
$2.00 of which is for a year’s subscription to Madrofio. Dues should be
remitted to the Treasurer. General correspondence and applications for
membership should be addressed to the Secretary.

wae

Complete Your Files! ‘

MADRONO- |
A West American Journal of Botany A

“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges. ae

Volume I, 1916-1929. . . $5.00 ny
Volume II, 1930-1934 .. 5.00 ?
Volume III, 1935-1936 . 5.00 by
Single numbers ....... 0.75 i

There is only a limited supply of Vol-
umes I and II. i

Address all communications u
and orders to: ia

Dr. David D. Keck : 4
Carnegie Institution of Washington la
Stanford University, California |

1938] COPELAND: ALLOTROPA | 137

THE STRUCTURE OF ALLOTROPA
Hersert F. Copetann

The monotropoid genus Allotropa, with the single species A.
virgata Torrey and Gray, was described by Gray (9) as “gathered
on the Cascade Mountains of northern Oregon, by Dr. Pickering
and Mr. Brackenridge, in the South-Sea Exploring Expedition
under Commodore Wilkes.” The “northern Oregon” of the time
of the Wilkes Expedition is
of course the present State of
Washington. The original
publication just cited em-
braced all the essentials of
taxonomic publication, the
description of a genus, the
name of a species, and the
citation of a specimen; but
it was brief and in a foot-
note; and a few years later
Gray (10) presented a more
extensive and formal ac-
count based largely on an
adequate collection by Bo-
lander, in the State Geologi-
cal Survey, “in Mendocino
County, between Little Bear
Harbor and Noyo, ‘generally
near Quercus densiflora.’ ”’
The plant is now known to
range from southern British
Columbia through Washing-
ton and Oregon and south in
California, in the Coast
Range to Sonoma County
and in the Sierra Nevada to

: Kings River. It occurs at
Fic. 1. Allotropa virgata. Plants col- ltideoe tf b
lected on Mount Shasta, California, by Nc UG re aa MO ated ONS
William Bridge Cooke, September 8, 1936. %€@ level up to eight or nine
Photograph by George C. Kimber. thousand feet. The habitat
of Bolander’s collection
caused Gray (11), and hence various European authors, to regard
oak forests as the typical habitat of the plant. Actually, our tan-
bark oak is currently excluded from oaks proper, as Lithocarpus
densiflora; and while Allotropa is often found as an associate of
this plant, it is perhaps more often found in coniferous forests.

Maprono, Vol. 4, pp. 137-168, January 11, 1938.

138 MADRONO [Vol. 4

Allotropa invites superficial attention as being infrequent and
at the same time conspicuous by its coloration in carmine and
white. More fundamentally, it invites attention by certain pecu-
liarities and uncertainties in its structure: authors have mentioned
and again denied the presence of bractlets on the pedicel; there
is a single whorl of perianth segments, interpreted sometimes as
petals, sometimes as sepals (Gray adopted the former interpreta-
tion in his first mention of the plant, and the latter in his second) ;
the anthers turn a half-somersault as the bud develops into the
flower, agreeing in this feature with those of Clethra and Pyrola
and differing from those of the other saprophytic Ericales; the
flower has been described as lacking a disk. Interpretation of
the characters affects, of course, opinion as to relationship. Gray
associated the plant first with Monotropa, then with Pyrola and
Schweinitzia, then with Pterospora. The more recent authorities,
Drude (8), Andres (1), Domin (7), Small (14), and Jepson
(13), have tended to separate it sharply from the other monotro-
poid plants, either implicitly by listing it first among the genera
of the group, or explicitly by assigning it to a separate group of
subtribal rank.

MaTERIALS AND MeEtTHODs

The anatomical part of the present study is based upon two
collections. Mr. Milo S. Baker of the Department of Botany,
Santa Rosa Junior College, California, was so kind as to send a
fresh specimen, collected about May 15, 1936, by Mr. McMillan
of Annapolis, Sonoma County. It consisted of a single shoot
without roots, in full anthesis. This collection is the basis of my
understanding of the flower. Later in the same year, I had the
good fortune to make the acquaintance of Mr. William Bridge
Cooke, who was botanizing on Mount Shasta during the summer
vacation. He kindly sent most abundant and excellent fresh
material collected on September 8. It included abundant roots
and nearly ripe fruits. My colleague, Mr. George C. Kimber,
had the kindness to photograph this collection (text fig. 1).

After preliminary study, the material as received was fixed
in Bouin’s fluid. It was imbedded and sectioned in standard
fashion. The staining combination generally used has been safra-
nin and light green; this makes conspicuous the xylem and other
tissue with lignified walls, and also cells containing stainable
material which I take to be tannin. In studying ovules and seeds,
in which lignified and tanniniferous cells are not of primary in-
terest, a combination of acetocarmine and aniline blue was found
useful. With the help and advice of Dr. A. S. Crafts and Dz
Katherine Esau, both of the College of Agriculture, University of
California, Davis, a method of seeing the phloem clearly was
worked out. The microtome sections are passed through xylol

1938 ] COPELAND: ALLOTROPA 139

and the alcohols down to water; exposed for some five minutes
to 5 per cent H,SO,; flooded alternately with 0.1 per cent aniline
blue and Crafts’ (6) killing solution (KI, 0.75 g.; I, 0.5 g.; H,O,
100 cc.), several changes of each, each for several minutes; and
mounted and examined in Crafts’ mounting solution (glycerine,
30) Cea O.g 00) ce.; ZnCl, +2) oon trace; I, 0.2 ¢.). Such
preparations are not permanent in the sense that they can be
stored away and studied in subsequent years; but they will last
for months, and even improve, at first, with age.

Tue Root System

As in other monotropoid plants, the root system is the perma-
nent member of the plant, the shoots being temporary reproduc-
tive structures. The collection studied was from sandy soil and
the roots were found fifteen to thirty centimeters below the sur-
face. Roots are of all sizes from less than one millimeter to
about four millimeters in diameter (pl.. XXII, fig. 2). The
smallest roots bear two rows of branches; the larger ones bear
four rows. I have been unable to follow any of the larger roots
to the tip; presumably they taper but little. In the microtome
sections, only one tip of a small root was studied. <A cap is pres-
ent; as seen, it is minute, consisting of shrivelled cells whose re-
mains are scattered along the sides of the root for a few milli-
meters back of the tip (pl. XXII, fig. 3). There is a definite
dermatogen. There are no root hairs, and no mycorrhiza was
detected. Spiral tracheids are formed to within a fraction of a
millimeter of the tip; but the plerome cannot be said to reach to
the dermatogen. The endodermis is not differentiated from the
cortical parenchyma; accordingly, the boundary between the

broad pericycle and the cortex cannot be recognized until con-
_ siderable development has taken place. The oldest part of a root
has a diarch primary xylem lying in a plane parallel to the axis
of the parent root. Branch roots originate in the pericycle oppo-
site the edges of the strand of xylem and grow out through the
cortex; the boundary between stele and cortex is first evident by
the level at which branch roots originate (pl. XXII, figs. 4, 5).
The xylem of the branch root becomes connected to the primary
xylem of the parent root. Apparently smaller roots remain
diarch; larger ones, diarch at first, begin to develop a tetrarch
xylem after forming their first branches, and subsequently form
branches in four rows. A cambium forms secondary xylem, most
abundantly in the angles between the four flanges of the primary
xylem (pl. XXII, fig. 6). The primary xylem consists chiefly of
scalariform elements opening into each other through scalari-
‘form or reticulate cross walls; with these are mixed many paren-
chymatous cells with non-lignified walls. The secondary xylem
is of similar conducting elements and includes vascular rays in

140 MADRONO [Vol. 4

some of which the cells are tanniniferous. As the secondary
wood appears, the line between stele and cortex becomes per-
ceptible by an indefinable difference between the cells inside and
outside a cylindrical boundary along which the walls of adjacent
cells become more or less stainable with safranin. At this level,
and in the planes where branch roots have already developed,
adventitious buds appear.

I am not clear as to the timing of the development of which
stages were seen. I suppose that growth in length and formation
of branch roots takes place in early summer; that formation of
secondary wood takes place next, and the development of buds
in late summer; and that the shoots appear above ground, flower,
and fruit, in the summer after their origin. Presumably each
root bears only one crop of secondary roots, and only one crop
of shoots, and then dies, leaving the space to the branch roots
which it has produced.

These roots, in contrast to those of Sarcodes and Pleuricospora
are perfectly normal; there is no question here of a procaulon.
The absence of mycorrhiza may be associated with the sandy soil
from which the material was collected (cf. Christoph (2)); and
the presence of a root cap and the endogenous origin of the
branches may in turn be associated with the absence of mycor-
rhiza.

Gross STRUCTURE OF THE SHOOT

Each stem is unbranched and can be regarded as a peduncle
together with the axis of a raceme. All of the foliar organs are
essentially of the same nature, although the lower ones may be
called leaves and the upper ones bracts in a raceme. When fresh
and living they are white and fleshy, acute, usually entire below
and more or less fimbriate above. They vary in size and shape
in different zones of the shoot. They show small numbers of
parallel veins evident as red streaks. They do not long remain
fresh and white, but soon shrivel and turn brown or black.

The most conspicuous feature of the plant is the coloration
of the stem. It is basically white but marked by a regular net-
work of brilliant red bands (pl. XXIV, fig. 21). The junctions
of this network lie at the insertions of the leaves; two lines con-
verging from below join at each node, ascend the stem a short
distance as a broad single line, and then diverge, running to the
bases of the two leaves next above.

In most of the published accounts the shoots are described
as rather short and divisible into a densely scaly lower part, a
sparsely scaly middle part, and a bracted raceme. Some plants,
however, are quite tall, and it is sometimes possible to distinguish
other regions in addition to the ones just mentioned: if the roots
are deeply seated, there may be a long slender stalk below the
densely scaly portion; and the latter is sometimes divisible into

1938 ] COPELAND: ALLOTROPA 141

a lower region of short leaves and an upper region of longer ones.
There is considerable variation in the relative sizes of these parts.
The phyllotaxy is spiral, but it is commonly not an orthodox
spiral: the numbers found by counting the files of scales on the
same shoot, respectively to right and to left, are often not ad-
jacent numbers of the 2, 8, 5, 8, ete., series; they are often not
numbers of this series at all. Tables I and II present some frag-

TABLE I
Lengths in centimeters of various parts of shoots of Allotropa

; : Stalk Region of
ete ne ODI low compact scales Peduncle Lint Total
of shoot ————— rescence
L U
ower pper
A he er 5 17 17 35
B 4 5 6 16 20 51
C 2.5 am 14 8.5 9 34
D 2 - 11 6 is 26
E 2.5 14 5 5 26.5
F 1.5 14 6 5 26.5
G 3 6 11 8 28
H 6 5 14 9 34
I 3 1.5 9.5 16 30
J 1.5 16 9 7 33.5
K 10 9.5 4.5 5 29
TABLE II

Numbers of parstichies to left (1) and right (7) in various parts of
certain shoots of Allotropa

‘ ; Region of
Designation compact scales Pedancle Inflo-
of shoot ate EN eee Ee ee ee rescence
Lower Upper

is 151 51 6 1
16r 6r a» Ar
101 bE Al 31
7 Vere 3r Ar

81

6r

Various other Al

shoots of 57
Cooke’s 31 21
collection 5r 5r
51 31
5r 5r

mentary notes on these points. In these tables, shoot “A” is the
one furnished by Mr. Baker; shoot “B” is the exceptionally tall
one of Mr. Cooke’s collection, shown in the middle of figure 1;
the others are various other shoots of the latter collection.

142 MADRONO [Vol. 4

ANATOMY OF THE SHOOT

By sectioning a fresh stem, one finds that the red pigment is
confined to the epidermis (in the ovary and stigma it is found
also in the next few layers of cells within). It is an indicator,
turning blue in ammonia vapor and red again in hydrochloric
acid vapor; evidently it is an anthocyanin. It disappears com-
pletely during fixation and imbedding. |

In microtome sections the epidermis both of stems and of
leaves is found to bear a thin cuticle with very fine lengthwise
striations; to be without stomata; and to contain tannin in every
cell

The cortex and pith are of parenchyma with many scattered
tanniniferous cells.

The vascular cylinder is a typical siphonostele, such as we
expect in woody plants rather than in herbs: it is broken by
vascular rays (bands of parenchyma, frequently tanniniferous)
and by:leaf gaps, but not by pith rays. At the very base of the
shoot, the vascular cylinder is compressed parallel to the axis of
the root which bears it, and is connected to the secondary wood
of the root.

The xylem includes little clusters of spiral tracheids, more or
less crushed as the stem matures, on the inner border; it consists
chiefly of scalariform vessels with scalariform or reticulate cross
walls. The phloem includes, beside much parenchyma, little
patches of typical sieve tubes and companion cells; there is usu-
ally one such cluster between each two vascular rays. There are
no more than traces of cambial activity (pl. XXIII, fig. 11).

In young shoots the pericycle outside the phloem is paren-
chymatous (pl. XXIII, fig. 8). Later, in shoots still underground
but well developed and ready for flowering in the following
season, one finds a heavy sheath of lignified fibers between the
phloem and the cortex (pl. XXIII, fig. 9). In fruiting shoots,
several layers of cells of the pith, not directly against the xylem
but separated from it by a narrow band of parenchyma, are ligni-
fied, and form a sheath on the inner side of the vascular cylinder
(pl. XXIII, fig. 10).

Each leaf trace consists of a single bundle leaving a gap in
the vascular cylinder. Branch traces vary in character with
height in the stem and stage of development; the variations
parallel those of the fibrous tissue already mentioned. No branch
traces are to be found in association with the lowest leaf traces
of the shoot. In older material, from higher in the stem, branch
traces exist as pairs of bundles running from the two sides of
each leaf trace and ending under abortive axillary buds. A
sheath, in the form of a trough-shaped band of fibers, lies against
the outer side of the leaf trace, and an inconspicuous little strand
of fibers accompanies each bundle of the branch trace (pl. XXIII,

1938 ] COPELAND: ALLOTROPA 143

x

weer e ee ew
=.

es
Se -

7--- --

Prate XXII. Arxrorropa vircata. Fig. 2. Roots and base of shoot X 1.6.
Fig. 3. Longitudinal section of root tip x 320. Figs. 4, 5, 6, '7. Cross sections of
roots X16; fig. 4, a small root with diarch primary xylem; fig. 5, a larger root
showing tetrarch xylem and attachment of a branch; fig. 6, root showing secon-
dary xylem and origin of a bud; fig. 7, root bearing a well developed bud.
Primary xylem and tanniniferous vascular rays black; secondary xylem heavily
stippled; phloem and precambial tissue lightly stippled.

144 MADRONO [Vol. 4

fig. 9). In the fruit-bearing rachis, leaf-trace and branch-trace
(that is, flower trace) are united into a cylinder of tissue com-
pletely enclosed by a sheath of fibers (pl. XXIII, fig. 10). This
body extends upward in the cortex through a distance of several
centimeters before reaching the level of the bract and flower

which it supplies (pl. XXIV, figs. 183-20. Note that in the models .

the vertical scale is only a quarter of the horizontal scale; these
traces are exceedingly long and slender). At the level of the
bract, the leaf trace escapes as several small bundles; here, in the
inflorescence, the leaf trace appears as a minor appendage of the
branch trace. The sheath may fade out at this level, or may con-
tinue into the pedicel; it fades out below the receptacle.

Tue FLOWER

The pedicel in flower is white, approximately 1 millimeter
thick and long. Referring to Bolander’s collection, Gray re-
marked that “two subulate bractlets which were thought to have
been detected upon the original specimen of Allotropa are not
seen in the good specimens now in hand.” Drude (8) and Domin
(7) both write of these bractlets as definitely present. The fact
is that they are present on some specimens, but absent from the
majority, at least of specimens collected in California.

The five perianth segments (less commonly four or six) are
white, more or less rhombic in outline, fimbriate, commonly a
shade under 5 millimeters long and slightly wider.

There are twice as many stamens (pl. XXV, figs. 25-30) as
perianth segments. The filaments are glabrous, dark, about as
long as the perianth segments. The summit of the young fila-
ment is bent outward at a right angle and attached to the base of
a dark anther about 2 millimeters long. This juvenile position is
approximately the morphologically normal position of the anther;
the outer side is the dorsal side. Even in the youngest material
available to me, the anthers are dehiscing; each one opens through
two lengthwise slits from the lower end half-way up the dorsal
side. The slit in each lobe crosses the ends of both microspo-
rangia of the lobe; the wall between them has already broken
down in the youngest material I have seen. As the flower ma-
tures, the angle at the summit of the filament changes, the end
swinging inward through ninety degrees, so that the anther hangs
downward with the dorsal side inward and the slits at the top.
If the anther is allowed to dry, the slits gape widely and appear
as pores. A vascular bundle passes up within the filament, bend-

ing as the filament does; makes an additional right angle as it —

enters the anther; and traverses the middle of it, in the septum
between the two lobes and in the plane of the septa between the
two microsporangia of each lobe.

1938 ] COPELAND: ALLOTROPA 145

nae at
IM hese

Pratt XXIII. Attorropa vircata. Fig. 8. Cross section of the base of a
young shoot x8. Fig. 9. Cross section of a more highly developed shoot x8.
Fig. 10. Cross section of rachis X 8. Xylem and external sheath black; phloem
and small-celled parenchyma within the xylem stippled; internal sheath cross-
hatched. Fig. 11. Details of the cross section of the vascular cylinder, from the
point indicated by the dotted square in fig. 9, x 320.

146 MADRONO [Vol. 4

The pollen grains are solitary and binucleate, in these re-
spects being typical of the monotropoid group. In the material
studied, most of them were found to be 8-grooved (pl. XXV, fig.
35); thus differing from those of Pterospora and Pleuricospora,
which are usually 4-grooved.

The ovary is bright red, subglobular, and usually about 5 milli-
meters in diameter. It bears at the base a whorl of inconspicu-
ous nectaries projecting between the bases of the filaments. It is
impressed at the top, so that there is a groove about the base of
the style, which is white, usually about 1 millimeter long and
thick. The stigma is red, ordinarily about 1 millimeter high and
3 millimeters in diameter, obscurely 5-lobed (varying to 4- or
6-lobed) ; grooves, one on top of each lobe, converge at the
center of the stigma on the opening leading to the interior of the
ovary. In fresh material, the stigmatic surface is covered with
a sticky fluid; microtome sections have shown germinating pollen
grains imbedded in this.

In internal structure the ovary (pl. XXV, fig. 24) agrees with
the other pyroloid and monotropoid plants which are said to
show axile placentation: a cross section near the base shows five
distinct locules, opposite the perianth segments, with a massive
placenta projecting from a central column into each. Higher up,
each placenta is split radially into two parts; the clefts between
them radiate from a vertical channel continuous with the style
channel. In this upper part, then, the placentation is really pa-
rietal, with the placentae thrust deeply into a single cavity.
(What one calls a placenta in the lower part of the ovary is, as
is usual in examples of axile placentation, the two edges of a
single carpel; in the upper part, as is usual in examples of parie-
tal placentation, it is the united edges of two adjacent carpels.)
These placentae of the upper part, continuations of the septa
between the cavities below, continue into the style as ridges pro-
jecting into the style channel. The lobes of the stigma stand
above the grooves between them; each lobe is the end of one
carpel.

The inner surface of the ovary wall is armored with a layer
of sclerenchymatous fibers as in Pterospora, Pyrola, and Chima-
phila, but not Newberrya, Pityopus, and Pleuricospora. The pres-
ence of this layer is associated with the dehiscence of a capsule,
its absence with the production of a berry.

The stigmatic surface is covered with an epidermis of col-
umnar cells, laterally in contact with one another, with the free
ends domed outward. This is apparently a less specialized sur-
face than those of Sarcodes, Pterospora, Pityopus, and Pleuricospora,
in which these cells become spindly needles not in lateral contact
with each other. These cells, and also several inner layers, but
not the cells farthest from the surface, are tanniniferous.

I

In

i

|

i

AT
\

ci

Linnomartyro00

sf

COPELAND: ALLOTROPA 147

oT
pets

|

Il

Ay Ny) WW

n

WOT TTT TATWN

|

er
ne

vi
i

e

iW

UY ULI

1mm x 10

x10 | Cross
Sections
<—

Imm xX 20

19 25

Pirate XXIV. Attorropa vircata. Fig. 12. Longitudinal section of vas-
cular cylinder, from a stem comparable with those represented in figs. 9 and 11,
X 320. Figs. 13-18. Portions of cross sections of rachis in succession upward
x8. Fig.19. Model of a portion of the vascular cylinder of the rachis, based on
figs. 13-18, not showing the sheath. Vertical scale <4; horizontal scale x 16.
Fig. 20. Similar model showing the sheath. Fig. 21. External view of the rachis

X 1.6.

148 MADRONO [Vol. 4

The vascular cylinder in the pedicel in my material, which is
without bractlets, shows no trace of bundles supplying them.
The first whorl of five bundles in the receptacle forks periclinally,
the lower branch running to a perianth segment, the upper to a
stamen. The association of the perianth segment bundles with
stamen bundles, and the location of the perianth segments oppo-
site locules of the ovary, are features of petals as distinguished
from sepals in the other monotropoid genera, and constitute proof
that the perianth segments of Allotropa are petals. A second
whorl of five bundles, alternating with the first, consists of sta-
men bundles. The remainder of the vascular supply of the flower
breaks up, not without irregularity, into ten bundles. Five of
these, lying in the planes of the petals, soon bend sharply out-
ward and ascend the ovary wall; these are carpel dorsals. At
the summit of the ovary, they bend down to the base of the style,
then continue up the style in the thin places between the internal
ridges; they end in the lobes of the stigma. The last whorl of
five bundles, being in fact pairs of carpel ventrals, enters the
central column of the ovary near the inner edges of the septa.
They bend, branch, and break up in furnishing the vascular sup-
ply of the placentae. Branches of them may or may not run out
through the septa to the ovary wall. In older flowers and in
fruits, indefinite small numbers of bundles springing from the
bases of the staminal bundles ascend the ovary wall for some dis-
tance between and parallel to the carpel dorsals. These corre-
spond to the supernumerary bundles which puzzled me in Pity-
opus. They are evidently of no phylogenetic significance, but are
developed in response to need, or by accident, in irregular posi-

tions (pl. XXV, fig. 86).

SEED AND FRvIT

Ovules ready for fertilization, and nearly ripe seeds, have
been seen; the stages of development have not.

The ovules are typical of the monotropoid group. An ovule
has an integument of two layers of cells; the nucellus is already

Pirate XXV. ALLOTROPA VIRGATA AND Prerospora. Fig. 22. Bud from an
underground shoot which would flower during the following summer x40. Fig.
23. Flower X4. Fig. 24. Dissection of a flower X8. Fig. 25. Two stamens
from a bud x8. Fig. 26. Diagram of the anther of such a stamen to show the
course of the bundle supplying it, the locus of the wall between microsporangia
of the same anther lobe, and the plane of the section shown in fig. 34. Fig. 27.
Similar diagram of a stamen in the mature position. Fig. 28. Mature stamen,
lateral view <8. Fig. 29. Mature stamen, view from within the flower x8.
Fig. 30. Mature stamen after dehiscence X8. Figs. 31, 32, 33, stamens of
Pterospora X 8, for comparison: fig. 31, ready for dehiscence; fig. 32, in process
of dehiscence, lateral view; fig. 33, in process of dehiscence, view from outside
the flower. Fig. 34. Cross section of anther at point of junction with filament
X40. Fig. 35. Pollen grain X800. Fig. 36. Model of the vascular tissue in the
receptacle approximately x 40.

1938 ] COPELAND: ALLOTROPA 149

Puate XXV. ALLOTROPA AND PreROSPORA.
page 148.)

(See explanation of figures on

150 . MADRONO [Vol. 4

absorbed; the embryo sac consists of the usual egg, two syner-
gids, endosperm mother cell, and three antipodal cells (pl. XXVI,
fig. 40). The seed as a whole (pl. XXVI, fig. 42) is about 0.8
millimeter long; it shows a swollen middle part and a tail at each
end; it is nearly white. Microtome sections (pl. XXVI, fig. 41)
show that the integument, excepting the outmost layer of cells,

tends to be absorbed or to collapse; that the endosperm has a

haustorium at each end (the exact structure of the haustoria is
not clear: they seem to consist of a few empty, very thick-walled
cells) ; the endosperm proper few-celled, the cells well filled with
granular material; the embryo, staining in the same fashion as
the endosperm, 2- or 8-celled. There is probably a suspensor
which is absorbed during development. The few cells of the
embryo are separated by oblique walls, as if there were a termi-
nal cell from which the others were successively cut off. I be-
lieve my understanding of the embryo of Pleuricospora as recently
published (5) was incorrect. The embryo of Pleuricospora is of
the same appearance as that of Allotropa, and presumably shows
the same manner of growth.

The ovary seems to grow but slightly in developing into a
fruit (pl. XXVI, figs. 37, 38). I have, indeed, some quite large
fruits, about 1 centimeter in diameter; these are borne on an
exceptionally large shoot, and are presumably developed from
exceptionally large flowers. I have not observed dehiscence in
nature. Fruits dried in the herbarium show two sets of lines of
weakness, dehiscence taking place along both. Those of one set
lie along the planes of the carpel dorsal bundles, and result in the
ordinary dehiscence of a loculicidal capsule, as in Pyrola, Chi-
maphila, and Pterospora. The other set forms a circumferential
line about the base of the style, as in Sarcodes.:

DIscussion

The indefiniteness of various characters of Allotropa—in the

relative development of the parts of the shoot; in the phyllotaxy;
in the presence or absence of bractlets; and in the numbers
of petals, stamens, and carpels—arouses speculation. One is
tempted to interpret it in anthropomorphic terms as a resultant
of a relaxation of discipline permitted by the dependent manner
of nutrition. No scientific explanation of this indefiniteness is
forthcoming: one is ignorant of the genetic and physiological
mechanisms involved in such behavior, and of the manner of im-
pact of natural selection such that these plants are free to vary
much more widely than most.

This indefiniteness of character is itself a character of the
whole monotropoid alliance. Variation in the nature of the calyx,
discussed by Domin (7), is a phase of it. In perhaps a majority
of the genera involved there is a perfectly definite calyx of four

1938 ] COPELAND: ALLOTROPA 151

Pratt XXVI. Attorropa vircata. Figs. 37, 38. Fruits x4. Fig. 39.
Summit view of a fruit in process of dehiscence (induced by drying and teasing
with needles). Fig. 40. Ovule 320. Fig. 31. Longitudinal section of nearly
mature seed X 320. Fig. 42. Whole seed x 80.

152 MADRONO [Vol. 4

or five sepals alternating with the petals. In other genera, Mono-
tropa, Hypopitys, Newberrya, and Pityopus, the number of sepals
is variable, and the sepals tend to intergrade with leaves, bracts,
or bractlets; buds and flowers may appear in the axils of the
sepals of Pityopus, as of ordinary leaves. In Allotropa the calyx
is completely suppressed, unless the bractlets, present on a minor-
ity of the specimens, are its representatives. As asepalous rather
than apetalous, Allotropa is much less an anomaly than it has ap-
peared to be. As I understand Domin’s thesis, he suggests that
the monotropid plants are primitively asepalous, and that a calyx
(or pseudocalyx) has gradually been evolved from other foliar
organs. I would be inclined to read the series in the reverse
direction, and to understand first a loss in definiteness of numbers
and sharp distinction of parts, and subsequently a loss of the
sepals.

The stamens, and particularly the anthers, of Allotropa show
resemblances in several directions; most obviously, with those of
Clethra, Pyrola, and Chimaphila. In their mature position, they
are the same as anthers of Arbutus and Arctostaphylos; but they
lack the paired tails of the anthers of these genera. Finally, it
is clear that anthers of Pterospora (pl. XXV, figs. 31-33) are
essentially the same, although differing in being fixed in a hori-
zontal position with the dorsal side upward. If Allotropa lacks
the tails which anthers of Arbutus and Arctostaphylos possess,
Pterospora has them. Pterospora shows another interesting fea-
ture: in addition to the slit along the proximal end of the anther,
another is developed at right angles to this, along the plane sepa-
rating the two microsporangia of each lobe. This gives a hint as
to the evolution of the anthers of Pleuricospora and Newberrya,
which open by lengthwise slits in this latter plane.

I have already pointed out resemblances in several directions
as shown by the gynoecium. At this point one may offer some
strictures on a recent paper by Hunt (12). This author assumes
the evolution of carpels eventually from dichotomously branched
thalli, and interprets them on this far-fetched basis. He recog-
nizes two types, a more primitive 3-lobed type, exemplified by
Pyrola (and other examples), and a more advanced 1-lobed type
exemplified by Clethra (and again by other examples). The sug-
gestion that the carpels of Pyrola and Clethra are of different
types, and that the former is the more primitive is scarcely tena-
ble. To me, it is clear from Hunt’s figures that Clethra has a
pistil comparable with those of other Ericales, but slightly more
primitive; a less complete union of carpels is manifest in the fact
that the single style passage usual in Ericales branches into three
channels in the stigma of Clethra. The real difference is that in
Clethra each lobe of the stigma stands at the end of a single car-
pel; in Pyrola, each lobe is compound, consisting of halves of the

1938 ] COPELAND: ALLOTROPA 153

ends of two adjacent carpels. This distinction is interesting:
Allotropa and Sarcodes show the more primitive condition as does
Clethra; the genera with fully parietal placentation, Pleuricospora
and Newberrya, show as does Pyrola the derived condition.

The characters of Allotropa, as I have come to know them,
seem to me to bind together to a surprising extent the genera of
the Pyrolaceae of Engler and Prantl (the allies both of Pyrola
and of Monotropa), a group so heterogeneous that their common
origin has been questioned. As an asepalous plant, Allotropa can-
not be recognized as the direct ancestor of the other Monotro-
poideae; but we can conceive it to be the living genus which has
in most features diverged least from the extinct common ancestor
of the group. The authors who have listed it first among the
Monotropoideae seem fully justified.

Sacramento Junior College,
Sacramento, California, June, 1937.

LITERATURE CITED

1. Anpres, H. Piroleen-Studien. Verh. bot. Vereins Prov. Brandenburg 56:
1-76. 1914.

2. Christoph, H. Untersuchungen iiber die mykotrophen Verhaltnisse der
‘“Ericales” und die Keimung der Pyrolaceae. Bot. Centralbl. Beih.
38’: 115-157, t. 8. 1921.

3. Copeland, H. F. The structure of the flower of Newberrya. Madrofio 2:
137-142. 1934.

4, ————————. On the genus Pityopus. Madrono 3: 154-168. 1935.

5. ————————. The reproductive structures of Pleuricospora. Madrono 4:
1-16. 1937.

6. Crarts, A. S. A technique for demonstrating plasmodesma. Stain tech-
nology 6: 127-129. 1931.

7. Domin, K. Vergleichende Studien iiber den Fichtenspargel. .. . Sitz-
ungsber. kgl. bédhm. Gesellsch. d. Wiss., II Cl., I Stiick: 1-111. 1915.

8. Drupr, O. Pirolaceae. in A. Engler und K. Prantl. Die natiirlichen
Pflanzenfamilien 4°: 3-11. 1889.

9. Gray, A., Joun Torrey, and J. S. Newserry. Exogenous plants. Rep.
Expl. Pac. R. R. 6°: 65-90. 1857.

10. Gray, A. Characters of new plants from California and elsewhere. .. .
Proc. Am. Acad. 7: 327-401. 1868.

11. ———————.. Synoptical Flora of North America, Vol. II, Part I. New
York. 1878.

12. Hunt, K.W. A study of the style and stigma, with reference to the nature
of the carpel. Amer. Journ. Bot. 24: 288-294. 1937.

13. Jerson, W. L. Manual of the flowering plants of California. Berkeley.
1925.

14. Smatyt, J. K. Monotropaceae. North American Flora 29: 11-18. 1914.

154 MADRONO [Vol. 4

AN ANOMALOUS NEW SPECIES OF LAPSANA
FROM CHINA

G. L. STessins, JR.

In going over the unidentified specimens of the Cichorieae in |
the University of California Herbarium, I found one, collected
by Dr. A. N. Steward of the University of Nanking, which was
strikingly different from any species known to me in the tribe,
and which did not fit in any of the genera listed by Hoffman in
Die Natiirlichen Pflanzenfamilien. Although the plant strongly
resembles the oriental species of Lapsana, as well as some species
of Iveris (Lactuca spp. of auth.), the pappus consists of stout
hooks, totally different from any of the numerous types of pap-
pus previously known in the tribe. I sent a few achenes from
this specimen to Dr. S. F. Blake of the U. S. Department of Agri-
culture, and to Dr. A. R. Horwood at Kew, England, both of
whom wrote back that they did not recognize them as resembling
any Cichoriaceous plant known to them. Dr. Blake noted a re-
semblance to “‘one or two genera in other tribes with a similar
stiff uncinate pappus, of which the most striking is Hypericophyl-
lum of the Tagetinae.”’

Upon request, Dr. Steward kindly furnished me with addi-
tional fragments from the original collection, as well as further
information on its habitat, which was the flood plain of the
Yangtze River, near Ta T’ung, Anhwei. After careful compari-
son of this material with specimens of Lapsana apogonoides Maxim.,
with which species Dr. Steward had identified it, I found that the
anomalous plant resembled this species not only in habit, but in
the character of the involucres, corollas, styles, anthers, etc., as
well as in the shape of the achenes. Furthermore, although the
achenes of L. apogonoides are characteristically epappose, as is
typical of the genus, they occasionally bear reduced hooks (pl.
XXVII, fig. E). Hence the specimens from Ta T’ung are evi-
dently an anomalous species of Lapsana, related to L. apogonoides.
It may be described as follows:

Lapsana uncinata Stebbins sp. nov. Herba glaberrima pa-
tenta; folia basalia runcinato-pinnatifida segmentis deltoideis
obtusis remote denticulatis, segmento terminale majore. Caules
plures decumbentes; folia caulini reducta vel nulla, lobis saepe
acutis. Capitula 5-15 pro inflorescentia; peduncula elongata,
erecta. Involucra ad anthesin 5.5 mm., ad fructificationem 6.5
mm. longa, phyllaria exteriora minuta, deltoidea, parce ciliata;
phyHaria interiora 5—6, ad fructificationem elliptica, apice obtusa;
floscula 10-12 pro capitula. Corollae flavae, 8-9 mm. longae,
tubo brevissimo; antherae nigrescento-virides, 2.8—3 mm. longae;
styli rami filiformi, 1-1.2 mm. longi. Achaenia compressa, 2.8—3
mm. longa, flavescentia, scabra, costis 10-14, 2—4 valde crassiori-

1938] STEBBINS: LAPSANA 155

W F WG

Puate XXVII. Figures A-D, Lapsana uncinata Stebbins: A, habit x 1/3;
B, corolla x 2 2/3; C, involucre X 2 2/3; D, achenes x5 1/3. Fig. E, Lapsana
apogonoides Maxim., achenes X5 1/3. Fig. F, Lapsana humilis (Thunb.)
Makino, achene 51/3. Fig. G, Lapsana communis L., achene 5 1/3.

156 MADRONO [Vol. 4

bus; pappus uncinis 2—4, 1.2-2.2 mm. longis constans (pl. XXVII,
figs. A-D).

Plant completely glabrous, spreading; basal leaves 4-10 em.
long, runcinate-pinnatifid, the lobes deltoid, obtuse, remotely
denticulate, the terminal larger. Stems several from the root
crown, 7-15 cm. long, decumbent, leafless or with a single, re-
duced cauline leaf, its lobes acute. Inflorescence of 5—15 heads
on slender, erect peduncles 1.5—5 cm. long, these subtended by
small, acute bracts. Involucres in flower about 5.5 mm. long,
elongating in fruit to about 6.5 mm., calyculate, the outer bracts
small, deltoid, remotely ciliate, the inner 5-6, lanceolate in flower,
becoming much broadened and more or less concave in fruit, ob-
tuse at the apex, the midrib obscure below, but often thickened
above into a cord-like protuberance. Florets 10-12 per involucre,
the corollas pale yellow, 8-9 mm. long, spreading at anthesis,
their tube 1 mm. long, glabrous. Anther tube greenish to black-
ish, 2.8-3 mm. long, style branches filiform, yellow, 1-1.2 mm.
long. Achenes compressed, 2.8—-3 mm. long, stramineous or red-
dish yellow, scabrous, 10-14 ribbed, 2 or 4 of the ribs much
stronger than the others, pappus of 2—4 stout hooks, these 1.2—2.2
mm. long and minutely scabrous.

Cuina. Near Ta T’ung, Anhwei, 22 April 1924, A. N. Steward
5248 (type Herb. Univ. Calif. no. 234,000, duplicate in Herb.
Univ. Nanking, China). Fragments from the type collection are
in the United States National Herbarium, the Gray Herbarium,
the herbaria of the New York Botanical Garden, and of the Royal
Botanic Gardens, Kew, England.

Although this species simulates Lapsana apogonoides Maxim.,
except in its peculiar pappus, a few other differences between the
two species may be noted. JL. apogonoides has much smaller
corollas (4-5 mm. long) which are hardly at all spreading at
anthesis, shorter anthers (1.5 mm. long), larger achenes (3.5—4.2
mm. long) which are paler in color, and less prominently ribbed.
L. humilis (Thunb.) Makino, which also has the habit of L. apo-
gonoides and L. uncinata, has still smaller flowers, and reddish
achenes which about equal in size those of L. uncinata, but are
somewhat more flattened and have thinner ribs (pl. XX VII, fig. F).

The achenes of these three species may be ranged in a series
showing, from Lapsana uncinata through L. apogonoides to L. humilis,
a progressively greater reduction of the pappus and the strength
of the ribs, along with an increasing degree of compression of the
achene. This is accompanied by an increasing reduction in the
size of the involucres and florets. This tendency toward a phylo-
genetic reduction in the size of the floral parts can be found in
most genera of the Cichorieae. It has been noted by Babcock
and Cameron in Crepis (Univ. Calif. Publ. Agr. Sci. 6: 287-324.
1934) and by Babcock and Stebbins in the related genus Youngia

1938 | MASON: LINANTHUS 157

(Publ. Carnegie Inst., Wash. no. 484. 1937). JL. uncinata, there-
fore, may be considered a primitive, relic species, representing a
type ancestral to the other oriental species of Lapsana.

Lapsana communis, with its European and western Asiatic rela-
tives, represents a different group of species, whose relationship
to the oriental species on the basis of leaf shape, the character of
the involucres, corollas, anthers and styles, and the general shape
of the achenes, is undoubtedly fairly close, but which differ in
their upright habit, the greater compression of their achenes, and
certain details such as the pubescence of the corolla tube and the
color of the anthers and the style branches. In their floral and
achenial characteristics these species may be compared with the
more reduced of the oriental species. Hence L. uncinata is in
floral characteristics the most primitive species of its genus, and
its type of pappus, probably the original one in Lapsana, has been
lost in the other species through reduction. Therefore if the
pappus is to be considered a primary criterion of classification in
the Cichorieae, as it has by most students of the tribe, Lapsana
must be considered to occupy a very isolated position in it. How-
ever, the resemblance of this genus, not only in habit but in in-
volucral and floral characteristics as well, to two oriental genera
of the Crepidinae, Iveris and Youngia, is considerable, and strongly
suggests an actual relationship between them.

University of California, Berkeley,
August, 1937.

TWO NEW SPECIES OF LINANTHUS FROM WESTERN
NORTH AMERICA

Hersert L. Mason

In the progress of studies in the genus Linanthus of the fam-
ily Polemoniaceae the following new species have been dis-
covered. It will be noted in the descriptions that the term ‘tube’
as applied to the corolla is used in a somewhat restricted sense.
The corolla in this genus varies from salverform through funnel-
form to campanulate. This variation is effected through differ-
ence in proportion of the tube, the throat and the lobes of the
corolla. The term ‘tube’ is herein used to apply only to the es-
sentially cylindrical portion of the corolla, whereas the term
‘throat’ designates that portion of the corolla expanding from the
top of the tube to the base of the corolla lobes. The proportion
of these three regions of the corolla to one another is often an
important diagnostic character of species. Likewise the position
of the stamens on the throat or on the tube is usually constant
within the species.

In citing specimens, the following abbreviations are used for
the various herbaria: University of California, Berkeley (UC),

158 MADRONO [Vol. 4

Utah State Agricultural College, Logan (IU), State College of
Washington, Pullman (WS), Willamette University, Salem, Ore-
gon (WU).

Linanthus mohavensis sp.nov. Herba annua compacta nana
1-6 cm. alta; cotyledones oblongae; folia opposita, supra pilosa,
subtus glabra, in tres lobos lineares acerosos palmatim divisa,
lobis interdum pro parte connatis valde venosis, venis usque ad
basin distinctis; flores nocturni; cymae dichotomae, floribus soli-
tariis termalibus; calycis tubus interdum ad basin versus sparse
pilosus infra sinus usque ad basin membranaceus post anthesin
accrescens demum intra nervos propter capsulae dehiscentiam
fissus; calycis lobi intus pilosi, aliter glabri, demum inaequales,
apicibus acerosis; corolla lutea vel faucibus interdum purpureo-
tinctis 5-7 mm. longa calycem aequans vel vix longior denique
capsula accrescente expulsa; stamina circa 1 mm. longa faucium
basi affixa; filamenta glabra; anthera orbicularia; stylus usque
ad medium in tres lobos lineares divisus; capsula cylindrica tri-
locularis tubum calycis sub aequans; semina in quoque loculo

plures reniformia apud hilum constricta sub aqua inmutata. (PI.

XXVIII, figs. k-p.)

Diminutive compact annual, 1-6 cm. high; cotyledons oblong,
leaves opposite, pilose above, glabrous beneath, with prominent
veins free to the base, palmately divided into 3 linear, some-
times partially united acerose lobes; anthesis nocturnal; inflores-
cence dichotomously cymose, flowers solitary in the forks of the
cyme; calyx membranous to the base below the sinuses, the mem-
brane growing with the calyx and at length splitting with the
dehiscing capsule, lobes pilose within, calyx otherwise glabrous
or with a few scattered hairs below, tips acerose, becoming un-
equal; corolla yellow, sometimes purple-tinted in the throat, 5-7
mm. long, equalling or barely exceeding the calyx, at length
pushed out by the growing capsule; stamens included, about 1
mm. long, inserted at the base of the throat of the corolla, fila-
ments glabrous, anthers orbicular; style included, divided to
about the middle into 3 linear lobes; capsule cylindrical, about
equalling the calyx tube, 3-celled; seeds several to each carpel,
reniform, strongly constricted at the hilum, not mucilaginous
when wetted.

West side of Searles Lake, one mile south of Trona, San
Bernardino County, California, March 29, 1935, H. L. Mason 8232

(type, U.C. Herb. 567794); Poison Canyon, San Bernardino

County, California, April 30, 1935, H. L. Mason 8294.

Linanthus mohavensis Mason occurs on gravelly talus and mesas
of the Searles Lake region, in association with Linanthus Jonesiz
(Gray) Greene, Gilia latifolia Gray, Mohavea brevifolia Cov., Pha-
celia pachyphylla Gray and Atrichoseris platyphylla Gray. Al-
though growing in association with Linanthus Jonesu Gray and

1938 ] MASON: LINANTHUS 159

being most closely related to it by virtue of several characters in
common, it does not even superficially resemble it in field aspect.
The low compact mode of growth, the smaller flowers, the stamen
insertion, the nature of the pubescence, and the forked leaves
readily distinguish it from that species. The following key will
serve to point out the relationships of the four species in this
section of the genus.

Testa of the seed bladdery, hyaline and membranous at
least on the angles; seed ellipsoid, depression of the
hilum not conspicuous; calyx glabrous; plants 5-30
em. high; corolla usually white with brownish-purple
patches on the back (occasionally yellow in no. 2).

Filaments with a hairy pad at the base; leaves

palmately “diviGeds janes aoe eee 1. L. dichotomus
Benth.
Filaments glabrous, leaves simple ............... 2. L. Bigeloott

(Gray) Greene
Testa not bladdery nor hyaline, closely investing the seed;
seed reniform or subreniform, deeply constricted at
the hilum; calyx pubescent; plants 2-10 cm. high;
corollas commonly yellow.

Leaves simple; calyx glandular-pubescent exter-
nally; stamens inserted in the corolla-tube ... 3. L. Jonesti
(Gray) Greene
Leaves 3-lobed; calyx glabrous externally; stamens
inserted at the base of the corolla-throat ..... 4, L. mohavensis
Mason

Linanthus septentrionalis sp.nov. Herba annua erecta cauli-
bus plerumque simplicibus, 5-30 cm. altis; folia opposita glabra
vel pubescentia segmentis linearibus palmatim 5-7-partita; cymae
paniculatae, pedicellis filiformibus 5-20 mm. longis; calycis tubus
infra sinus usque ad basin membranaceus, lobis fere ad apices
membranaceo-marginatis, cum calyce pari passu accrescentibus ;
corolla 1-4 mm, longa, calyce sesquilongior, lobis albis vel pal-
lide coeruleis patente rotatis campanulatisve, faucibus tubo brev-
ioribus, annulo piloso media parte vel rarius basi filamentorum
instructis; stamina aequalia exserta basi faucium affixa; filamentis
glabris vel basi sparse pilosis; stylus exsertus; stigma trilobum;
capsula cylindrica trilocularis; semina in quoque loculo 2-4 sub
aqua mucilaginosa. (Pl. XXVIII, figs. a-e.)

Erect annual, usually simple, 5-30 cm. high; leaves opposite,
palmately divided into 5—7 linear segments, glabrous or pubes-
cent, 5-20 mm. long; inflorescence cymose-paniculate; flowers
solitary on filiform pedicels, 5-20 mm. long; calyx membranous
to the base below the sinuses, the membrane flanking the lobes
almost to the tips above the calyx-tube, growing with the calyx;
corolla 114 times the calyx, 1-4 mm. long, the lobes rotately or
campanulately spreading, the throat shorter than the tube, with
a hairy ring on the middle portion above the point of stamen-

160 MADRONO [Vol. 4

insertion or more rarely only on the base of the filaments; lobes
white or pale blue; stamens inserted at the base of the throat,
equal in length, exserted, filaments glabrous or with a few hairs
at the base; style exserted, stigma 3-lobed; capsule cylindric,
8-celled, each cell with 2-4 seeds; seeds producing mucilage when
wetted.

Camp Roosevelt, Tower Junction, Yellowstone National Park,
Wyoming, July 10, 1936, H. L. Mason 3497 (type, U.C. Herb. .
552425).

Linanthus septentrionalis Mason is a small flowered delicate
annual of the Great Basin and Rocky Mountain region which
superficially, closely resembles L. Harknessa Curran of the high
Sierra Nevada, the Cascade Mountains and westward. In general
it is more robust, has a larger corolla, with hairs on the inside as
well as sometimes on the filaments and has many more seeds to
the capsule than does L. Harknessui. These characters however
place it in close relationship with LZ. pharnaceoides (Benth.)
Greene rather than with L. Harknessu. The following key will
bring out the characters that may serve to separate the three
Species.

Seeds 1 to each cell of the ovary; corolla glabrous within,
barely exserted from the calyx; filament glabrous .. 1. LZ. Harknessii
Curran
Seeds several to each cell of the ovary; corolla rarely
glabrous within, usually with a hairy ring at or above
the stamen insertion, or the base of the filaments
hairy.

Corolla 2-4 mm. long, 114-2 times the calyx; hairs
dense to sparse in a ring on the throat of the
corolla, rarely absent; filaments glabrous or
with a few hairs; plants from east of the
Sierra and Cascade axis to the Rocky Moun-
GAINS Gr cereals c, V MR ee e ee 2. L. septentrionalis
Mason
Corolla 6-10 mm. long, 3-5 times the calyx; hairs
dense, on the base of the filaments only;
plants from the northern Great Basin and
the valleys and foothills of California ...... 3. L. pharnaceoides
(Benth.) Greene

EXPLANATION OF THE Figures. Puiate XXVIII.

Pirate XXVIII. Figs. a-e, Linanthus septentrionalis Mason: a flower;
b habit; c opened corolla; d seeds; e capsule and calyx. Figs. f—j, Linanthus
Harknessii Curran: f habit; g flower; h seeds; i capsule and calyx; j opened
corolla. Figs. k—-p, Linanthus mohavensis Mason: k habit; 1 flower; m capsule
and calyx; n seeds; o opened corolla; p leaf. Figs. q—u, Linanthus Jonesi
(Gray) Greene: gq capsule and calyx; r habit; s flower; ¢ seeds; wu opened
corolla.

1938] MASON: LINANTHUS 161

Pratt XXVIII. Comparative Sketcues oF LINaAntTHUus. (See explanation

of figures on page 160.)

162 MADRONO [Vol. 4

Specimens examined. British Cotumsia. Nicola, 8 mile creek,
6000 ft. G. V. Copley 68 (WS); Sophia Mountain, J. M. Macoun
66,561 (WS) ; Lake Osoyoos, J. M. Macoun 68,708 (WS). Wasu-
IncTon. Columbia County: Blue Mountains, C. V. Piper 2398
(WS). Lincoln County: Sprague, 1800 feet, Sandberg and Lei-
berg 201 (WS, UC). Okanogan County: east of Omak, C. B.
Fiker 698 (WS). Whitman County: Kamiak Butte, A. D. E.
Elmer 803 (WS). Oregon. Deschutes County: 10 miles east of
Bend, M. E. Peck 19,742 (UC, WU). Harney County: east side
of Harney Valley, J. B. Leiberg 2376 (UC). Ivano. Blaine
County: Camp Creek, Macbride and Payson 2956 (UC); Tikura,
Nelson and Macbride 1291 (UC, WS). Boundary County: Snowy
top Mountain, F. A. Warren 311 (WS). Kootenai County: Dry
Prairies, J. H. Sandberg 7219 (WS). Nez Perce County: 3500
feet, A. A. Heller 3482 (WS). Owyhee County: Silver City, 7000
feet, J. F. Macbride 358 (WS). Shoshone County: Clarkia, along
St. Maries River, C. R. Quick 1095 (UC). Montana. Carbon
County: Bridger, Bracket Creek divide, July 16, 1902, J. W.
Blankinship (WS). Gallatin County: Bozeman, J. W. Blankin-
ship 357a (WS); Spanish Basin, 6500 feet, Rydberg and Bessey
4821 (WS); 15 miles south of West Yellowstone, B. Maguire
1199 (IU). Jefferson County: Boulder Divide, Aug. 4, 1898, E.
N. Brandegee (UC). Mineral County: Bitterroot Mountains, H.
L. Mason 10,050 (UC). Missoula County: Fort Missoula, J. E.
Kirkwood 1294 (UC). Park County: Big Mud Creek, W. N. Suks-
dorf 376 (WS, UC). Powell County: Ovando, J. EL. Kirkwood
1424 (UC). Wyomine. Lincoln County: Headwaters of Hoback
River, 9000 feet, L. Williams 1281 (IU). Teton County: Snake
River at mouth of Pacific Creek, 6500 feet, L. Williams 1636
(WS). Yellowstone National Park: Camp Roosevelt, Tower
Junction, H. L. Mason 8497 (type, UC). Uran. Cache County:
Intervale, Blacksmith Fork Canyon, 5700 feet, B. Maguire 3670
(IU); Logan Canyon 20 miles above Logan, H. L. Mason 10,039
(UC); Tony Grove, J. B. Wann 3669 (IU). Wasatch County:
American Fork Canyon 8000 feet, 1902, M. FE. Jones (IU). NE-
vapA. Esmeralda County: W. H. Shockley 660 (UC). Nye
County: Toyabe Mountains north of Twin Lakes, Linsdale 758
(UC). Carirornia. Inyo County: White Mountains, V. Duran
2756 (UC). Mono County: Conway Grade, between Mono Lake
and Bridgeport, 7800 feet, D. D. Keck 2914 (UC).

_ University of California,
Berkeley, October, 1937.

1938 | GANDER: SAN DIEGO COUNTY ENDEMICS 163

NOTES ON SOME SAN DIEGO COUNTY ENDEMICS

Frank F. GAnpDER

Since the ranges of certain endemic species of San Diego
County are not well known it seems advisable to publish addi-
tional information furnished by specimens in the Herbarium of
the San Diego Society of Natural History. This is not a complete
list of all species endemic to this county. The collections cited
below help to define ranges or to extend them beyond reported
limits. Numbers cited in parentheses refer to herbarium acces-
sion numbers. Unless otherwise stated collections were made by
the author. I am indebted to Dr. Philip A. Munz of Pomona Col-
lege for some of the determinations.

Bioomeria CLEVELANDII Wats. Common on mesas and slopes
about San Diego, northeast to the vicinity of Fletcher Hills and
southeast to Otay Mountain: San Diego (10,992, 15,010), Daniel
Cleveland (6896, 6400, 6405, 6406, 6408); Woodwardia Canyon
of Otay Mountain (15,007); mesa northeast of Murray Dam,
near Fletcher Hills (15,088).

Bropiaca Orcutti (Greene) Baker. Occurs near streams, in
seepage areas, and in beds of vernal pools from San Marcos to
Otay Mountain: Kearney Mesa (11,099, 11,333) ; San Diego (15,-
216); four miles west of San Marcos (16,052); Woodwardia
Canyon of Otay Mountain (16,055).

Catocnortus Dunnu Purdy. Dry stony slopes of the Cuya-
maca Mountains at altitudes from 3500 to 4500 feet: Inspiration
Point, Florence Youngberg (11,740) ; Otay Mountain (11,722).

CHORIZANTHE OrcuTTIANA Parry. Rare; reported only from
Point Loma and Kearney Mesa: Point Loma, T. S. Brandegee
(1218), D. Cleveland § Charles Orcutt (7857); Daniel Cleveland
(7858) ;, Kearney Mesa (10,604).

CAULANTHUS STENOCARPUS Payson. Harbison Canyon south-
west of Alpine (14,699). Heretofore reported only from Ber-
nardo near Lake Hodges.

SepuM variecatum Wats. This yellow-flowered form is com-
mon in depressions of the mesas from Kearney Mesa to Otay,
and inland to El Cajon: East San Diego, Ralph W. Sumner
(2071); National Ranch, Daniel Cleveland (8324, 8325); El
Cajon, Daniel Cleveland (8326); near mouth of Mission Gorge
(15,954) ; San Diego State College (15,955) ; Lemon Grove (15,-
956); Otay (15,957) ; Kearney Mesa (16,031).

HEUCHERA BREVISTAMINEA Wiggins. Known only from the
Laguna Mountains: Shrine Camp, Vallecitos View, Charles F.
Harbison (12,088); Garnet Peak, Charles F. Harbison (12,089).

164 MADRONO [Vol. 4

RiBES CANTHARIFORME Wiggins. Common in the shade of
large boulders on sheltered slopes near Morena Dam and on the
north slope of Lyons Peak. On March 12, 1986, the writer
found many plants in full bloom on the sheltered hillside below
the dam (13,919). Every bush observed at this site was crowded
close against the shaded side of a large boulder and was often
completely hidden by the rest of the chaparral. At the type
locality, a short distance east of the south end of the Morena
Dam, one bush was found (13,918). On March 29, 19386, the
species was also found in abundance on the upper part of the
north slope of Lyons Peak, about 13 miles due west of the type
locality, extending from near the top, at 3755 feet, down to about
3100 feet (14,296). Most of the plants found here were in the
shelter of large boulders, but a few were seen in dense chaparral
in sheltered places. This species of Ribes may occur quite widely
in the southern part of San Diego County, as the area has been
incompletely explored botanically and it is easily overlooked.

ASTRAGALUS oocaRPus Gray. Occurs on dry slopes in our
mountains: Julian, Daniel Cleveland (8536); falls of San Diego
River, Daniel Cleveland (8537); Witch Creek (11,943); Santa
Ysabel (16,168, 16,201); Pine Hills (16,205); Carrizo Creek
near Lake Henshaw (16,171); Black Canyon near Mesa Grande
(16,332); Palomar Mountain (16,328). ~

CEANOTHUS cyANEUs Eastwood. Occurs in scattered clumps
in the chaparral from Mussey Grade to near Alpine, and abun-
dantly on the northeast slope of El Cajon Mountain: Mussey
Grade (15,098); four miles west of Alpine (15,279, 16,297) ;
Barona Valley (15,585); northeast slope of El Cajon Mountain,
altitude 3000 feet (15,628); southeast base of El Capitan (15,-
752); top of rocky bluff between San Vicente Creek and San
Diego River at their junction near Lakeside, the type locality,
M. Phillbrook & F. Gander (15,224).

CEANOTHUS AUSTROMONTANUs Abrams. Occurs in the chaparral
from near Cuyamaca Lake to near Lake Henshaw: Inspiration
Point (11,304); Pine Hills (16,111); Cuyamaca Lake (16,208) ;
south of Lake Henshaw on road to Mesa Grande.

Cruarxia DELIcATA (Abrams) Nelson & Macbride. Occasional
in openings in the chaparral near Mesa Grande and San Felipe
Creek to Campo, Potrero Grade and Harbison Canyon: five miles
south of Mesa Grande (11,359); burn in Harbison Canyon
(14,698).

ACANTHOMINTHA ILIctFot1a Gray. In adobe lands from Kear-
ney Mesa, San Diego, and Bonita inland to Viejas Grade: Viejas
Grade, Daniel Cleveland (6668, 6670, 6671); San Diego, Daniel
Cleveland (6669); Bonita, H. E. Hansen (11,457); San Diego

1938 ] GANDER: SAN DIEGO COUNTY ENDEMICS 165

State College (15,023); slope at mouth of Mahogany Canyon,
San Diego (15,451); slope at junction of Alvarado Canyon and
Mission Valley (15,453) ; Kearney Mesa (16,020).

Satvia Patmert (Gray) Greene. Apparently a hybrid be-
tween Salvia apiana Jepson and Salvia Clevelandi (Gray) Greene,
this species has been seldom collected: east of Mount Woodson,
near Ramona (16,064). The type material was secured by Dr.
Edward Palmer on the Tighe Ranch at the foot of the grade on
the old road between Ramona and Ballena. It has also been
reported from Torrey Pines Park and from Descanso Grade.

PoGoGYNE NupiIuscuLA Gray. Found only in depressions on
the mesas, mostly near San Diego: San Diego, (11,338), Mary S.
Snyder (4351); Daniel Cleveland (6672, 6675, 6676, 6677) ; Loma
Alta on Otay Mesa (11,557).

Pocoayne Azsramsu J. T. Howell. Known only from depres-
sions in the mesas near San Diego: San Diego (15,036), Daniel
Cleveland (6673) ; Miramar (11,332); Kearney Mesa (16,006).

MonarDELLA LANATA Abrams. Known previously only from
the region between Descanso and Alpine: Otay Mountain (11,-
679); Mt. Woodson (16,065, 16,070).

AmpBrosia PuMILA (Nutt.) Gray. Common around San Diego
and in El Cajon Valley, ranging north to San Luis Rey Valley,
south to National City, and inland to two miles east of El Cajon:
San Diego, Daniel Cleveland (86388, 8689, 8640, 8641, 8642); Old
Mission Dam in Mission Gorge (5647) ; Santee (11,492, 15,779) ;
east of El Cajon (12,059) ; Bostonia cut-off near railroad cross-
ing (15,777); junction of Bostonia cut-off and Lakeside road
(15,778); National City (15,581); Sweetwater Dam (15,999) ;
Bostonia (16,001) ; Lake Hodges (16,019) ; Mission Valley, west
of San Diego Mission (16,048).

GeRAEA viscipA (Gray) Blake. Known only from dry slopes
and hills in southern San Diego County: Campo, Fidella G. Wood-
cock (5569, 5570, 5571, 5572, 5578); George Stenberg (11,454) ;
Jacumba, Daniel Cleveland (8849) ; Hipass (15,295).

ARTEMISIA Patmert Gray. This species has been reported
previously only from Jamul Valley, National City, and La Jolla:
Jamul, Susan G. Stokes (13,463); east of Mount Woodson (16,-
066) San Vicente Creek near Foster (16,202); Bostonia cut-off
near junction with Lakeside road (16,338); Japatul Valley
(16,339).

Natural History Museum,

San Diego, California,
November 11, 1936.

166 MADRONO | [Vol. 4

THE CALIFORNIA NUTMEG TREE IN CULTIVATION

ALBERT WILSON

The interest in native plants is ever present among California
garden lovers. Catering to this interest are several nurseries
devoted entirely or in part to the culture and sale of native
plants. Such well known shrubs as Fremontia, Ceanothus, Car-
penteria, Romneya, Ribes sanguineum and Mahonia are often to be
found thriving
under (culti—
vation. Also
sprinkled in the
gardens of the
state are many
specimens of
California
pines, cypress,
sequoias and
even oaks.

Among the
coniferous na-
tives’ ote the
state, Torreya
californica Torr.,
a handsome tree
known com-
monly as Cali-
fornia nutmeg,
is; Lounids im va
few well estab-
lished gardens.
It is not unusual
to find trees of
great diameter
and height in
their native
habitat, but to
find a large Tor-

ee pas garden Fic. 1. Torreya californica growing at corner of the
PEASE ES) 0{E 18) old Seale estate, California Avenue and Byron streets,
ence. Of culti- Palo Alto, California.
vated speci-
mens the most outstanding is in the park opposite the post office
in Watsonville, California. Like a forest tree, singled out and
favored with an abundance of sunlight and moisture, this beauti-
ful tree is fifty feet high with a trunk three feet in diameter.

On the San Francisco peninsula, particularly in the Palo Alto
district, several noteworthy specimens of Torreya californica are

1938] REVIEW 167

growing as garden subjects. By far the most conspicuous is a
large tree in the Seale Tract on California Avenue at Byron
Street, Palo Alto. This tree tapers to a point and is forty feet
tall. The trunk is six feet in diameter at ground level and four
feet in diameter four feet above the ground. Here the trunk
leads off into many large branches causing the plant to appear
bushy, thus resembling an English yew. The branches spread
to a diameter of more than fifty feet. This tree was growing ad-
jacent to the old Seale mansion, which has just been torn down.
It might be safe to estimate its age as approximately sixty or
seventy years, since the Seale estate is about one hundred years
old, and all members of the family were garden minded.

In Menlo Park on the campus of the Park Military School for
Boys on Ravenswood Avenue, opposite the Southern Pacific
depot, there is another large cultivated specimen of the Cali-
fornia nutmeg. The tree, easily recognized by its dark foliage,
stands thirty feet tall, has a spread of twenty-seven feet and
arches decidedly to the west. The trunk is three feet in diam-
eter at ground level. Roughly estimated this tree is about fifty
years old. The garden, that of the old Mills estate, is of long
cultivation. On the grounds of the Leroy Nickel estate on Middle-
field Road in Menlo Park there is to be seen another fine speci-
men of Torreya californica. This tree is about thirty-five feet tall
with a three foot trunk at ground level and with a spread of
twenty-five feet to its branches. Mrs. Nickel reports that it was
planted about thirty years ago. A fourth Menlo Park specimen,

no doubt much younger than the others, is to be found on the
Donohoe estate on Middlefield Road. This tree is eighteen feet
tall.

Torreya californica may be regarded as very satisfactory in

_ cultivation and is to be recommended for the garden. All of the
_ specimens mentioned above produce abundant fruit and the
seeds germinate easily.

| Menlo Park, California,
March 13, 1937.

REVIEW: TWO NEW CATALOGUES OF CASCADIAN
PLANTS

The Flora of Crater Lake National Park. By F. Lytr Wynp.
The American Midland Naturalist. Volume XVII, Number 6.
Pp. 881-949. The University Press, Notre Dame, Indiana. 19386.

The Plants of Mt. Rainier National Park, Washington. By Haroip
St. Joun anp Frep A. Warren. The American Midland Natural-
ist. Volume XVIII, Number 6. Pp. 952-985. The University
Press, Notre Dame, Indiana. 19387.

The first is a very incomplete list, with keys, taxonomic notes
and a synonymy of the names used in the principal manuals, of

168 MADRONO [Vol. 4

the plants of the Crater Lake area. Neither habitat data nor
localities are given, but most of the specimens upon which the
list is based are in the Herbarium of the University of Oregon, at
Eugene. The other paper offers a few introductory remarks upon
the climate, topography, geology and vegetation of Mt. Rainier,
but the bulk of the work is a list of indigenous and adventive
species, without keys, but with habitat notes, some localities and
a few citations of specimens. Mr. Warren’s collection of speci-
mens, deposited at the State College of Washington at Pullman,
which substantiated many of the species included in this list, was
unfortunately destroyed by herbarium beetles several years ago.

It is a happy circumstance that the one treatment deals with
an important section of the southern Cascade Range, whereas the
other treats the highest and one of the northernmost peaks of
this range. The publication of any key or list of plants of the
Cascade Mountains is a welcome addition to the botany of the
Northwest and serves to call attention to the desirability of more
thorough studies of the same areas.—L. C.

NOTES AND NEWS

Word has been received from Honolulu, Hawaii, of the death
on November 7, 1987, of Mr. Leo D. Whitney, formerly a gradu-
ate student in the Department of Botany, University of Cali-
fornia, Berkeley. Mr. Whitney went to the Hawaiian Islands in
October, 1935, as Assistant Agronomist, Hawaii Agricultural
Experiment Station, and later was also appointed instructor in
the University of Hawaii. He was an enthusiastic and energetic
young botanist. News of his untimely death came as a shock to
friends here and in the islands.

With the objective of collecting certain fern species from the
type localities, Dr. and Mrs. E. B. Copeland left Berkeley, Cali-
fornia, on December 18, 1987, for Mexico. Expeditions will be
made to Acapulca, Oaxaca, and Orizaba.

Ynes Mexia left Berkeley, California, October 3, 1937, on her
fourth collecting trip in Mexico. At present she is at Balsas,
State of Guerrero.

Recently appointed research associates in the Department of
Botany, University of California, Berkeley, are: Dr. Mildred E.
Mathias, specialist in Umbelliferae; Dr. E. B. Copeland, recently
technical adviser and botanist, Bureau of Plant Industry, Philip-
pine Islands; Dr. F. W. Foxworthy, forest research officer, retired,
Federated Malay States.

hod

f

MADRONO ©

A West American Journal of Botany

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and stimulating articles dealing with
plant morphology, physiology, taxonomy,
and botanical history. These volumes
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brary and should be made accessible to
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Volume I, 1916-1929. . . $5.00
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NUMBER 6

-MADRONO

A WEST AMERICAN JOURNAL OF
BOTANY

C4

Contents

~ .PuHyLocenetic RELATIONS oF PINUS JEFFREYI AND Pinus ponperosa, NV. T.

AILEY DGS AT) RA NL AU aL OM eee hel! US a a Ae RT 169

On THE IDENTITY OF CAYTONIA NEVADENSIS Watson, Carl W. Sharsmith .. 171
Picea BREWERIANA in Swasta County, Philip G. Haddock............... 176
Tue Inwrropuction or Acacias Into Catirornia, H. M. Butterfield ....... 177
Harry Stantey Yates, Herbert L. Mason ................ 000.0. ee 187

_ Erioconum ABERTIANUM AND its Varieties, F. Raymond Fosberg ........ 189
AN UNDESCRIBED SPECIES OF Vi0LA FRoM Utanu, Milo S. Baker ........... 194

Reviews: Graham, Botanical Studies in the Uinta Basin of Utah and Colo-
rado (Mildred Mathias); reprint of Lindley, Illustrated Dictionary
of Botanical Terms (Mildred Mathias); Epling, The California Sal-
vias. A Review of Salvia, Section Audibertia (Ethel Crum); Presnall
and Patraw, Plants of Zion National Park (Herbert L. Mason) ;
Mathias, 4 Revision of the Genus Lomatium (Lincoln Constance) ;
Tischler, Die Bedeutung der Polyploidie fiir die Verbreitung der An-
giospermen, erldutert an den Arten Schleswig-Holsteins, mit Aus-

blicken auf andere Florengebiets (G. L. Stebbins, Jr.) .............. 196
Notes anp News: Ceanothus thyrsiflorus: Extension of Range (H. E.

PCIE eMC TOS TI ei wba Ma ae LAMM Mama oa Wi CuPc) a 199

OCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY ..................... 199

Published at Lime and Green Streets, Lancaster,
Pennsylvania

- April, 1938

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asprams, Stanford University, California.

Dr. Lincotn Constance, University of California, Berkeley.

Dr. H. F. Copetanp, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Business Manager—Dr. Davm D. Krecx
Lime and Green Sts., Lancaster, Pennsylvania
or
Carnegie Institution of Washington
Stanford University, California

Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

Established 1916. Published quarterly. Subscription Price $2.50 per year.
Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17,
complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num-
bers $0.75.

Papers up to 15 or 20 pages are acceptable. Longer contributions may
be accepted if the excess costs of: printing and illustration are borne by the
contributor. Range extensions and similar notes will be published in con-
densed form with a suitable title under the general heading “Notes and News.”
Articles may be submitted to any member of the editorial board. Manuscripts
may be included in the forthcoming issue provided that the contributor pay the
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any article are furnished at a cost of 4 pages, 50 copies $3.70; 100 copies $4.10;
additional 100’s .75¢; 8 pages, 50 copies $5.40; 100 copies $6.00, additional 100’s
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should be ordered when page proofs are returned.

Published at Lime and Green Streets, Lancaster,
Pennsylvania, for the

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Professor H. E. McMinn, Mills College, California. First Vice-
President: Mr. Maunsell Van Rensselaer, Blaksley Botanic Garden, Santa
Barbara, California. Second Vice-President: Mrs. Viola Brainerd Baird,
Berkeley, California. Treasurer, Dr. David D. Keck, Carnegie Institution of
Washington, Stanford University, California. Secretary: Miss Ethel Crum,
4004 Life Sciences Building, University of California, Berkeley.

Annual membership dues of the California Botanincal Society are $2.50,
$2.00 of which is for a year’s subscription to Madrofio. Dues should be
remitted to the Treasurer. General correspondence and applications for
membership should be addressed to the Secretary.

———

PSs =
a

ral
‘

Pe

1938] MIROV: PHYLOGENY IN PINUS 169

PHYLOGENETIC RELATIONS OF PINUS JEFFREYI AND
PINUS PONDEROSA

N. T. Mirov

\

At present the phylogenetic relations of Pinus Jeffreyi Balf.,
Jeffrey pine, and Pinus ponderosa Laws., ponderosa pine (west-
ern yellow pine), are not accurately known.

Although fossil three-needle pine material occurs as far back
as Jurassic, there is none of it which has been definitely assigned
to either of the species under consideration. These species may
be said to have practically no established fossil record, and it is
not certain which of the two pines is geologically older. Long
ago Lemmon (3) stated that P. Jeffreyi was not only an older
form than P. ponderosa, but that it should be considered the
ancestor of this comparatively younger species. Unfortunately
Lemmon does not give any support for his statement. Later

Simonsen and Rau (6) advanced a theory that pine species con-

taining in their oleoresin saturated fatty hydrocarbons are proba-
bly geologically older than the pines containing unsaturated
hydrocarbons, terpenes. It has been found by several investi-
gators that oleoresin of P. Jeffreyz contains a saturated hydro-
carbon, heptane, and no terpenes, while oleoresin of P. ponderosa
contains a mixture of terpenes and no heptane. Additional in-
formation concerning the differences in oleoresin composition of
the two pines may be found in a previous article by the author (5).

It has been suggested by Dr. Herbert L. Mason of the De-
partment of Botany, University of California, Berkeley, that,
from a distributional point of view also P. Jeffreyi is geologically
older than P. ponderosa. It is an endemic species restricted
almost entirely to California and associated with many other
endemic plants. On the contrary, P. ponderosa has a much larger
range that stretches far beyond the boundaries of the Jeffrey
pine endemic group. Hence, it is concluded that P. ponderosa is
the younger species. Moreover, Jeffrey pine is extremely stable,
that is, it does not vary much, thus apparently exhibiting char-
acteristics of racial senility. On the other hand, the extreme
variability of P. ponderosa may be considered an indication of its
relatively younger age.

The writer has been interested for some time in the chemical
composition of seed oil of the two species. It has been found
that the degree of unsaturation of the seed oil is much higher in
P. ponderosa than in P. Jeffreyi. In general, oil of pine seed con-
sists of triglycerides of unsaturated (oleic, linoleic and linolenic)
acids, the amount of saturated compounds being very small. The
degree of unsaturation of an oil is governed by the number of

double bonds in its molecules. Oleic acid has one, linoleic two,

Manrono, Vol. 4, pp. 169-200, April 15, 1938.

170 MADRONO [Vol. 4

and linolenic three, double bonds. With iodine an addition
reaction takes place; one molecule of oleic acid reacts with two
molecules of iodine, one molecule of linoleic with four, and one |
molecule of linolenic acid with six molecules of iodine. Iodine
number is simply the percent of iodine absorbed by the oil. An-
alysis of oil samples extracted from seed of the two pines has
shown that the iodine number of P. ponderosa varies between 147
and 154, the average of twenty samples being 151, while that of
P. Jeffreyi is between 129 and 1388, with 134 as an average for
twenty samples. In the following discussion the writer will at-
tempt to show how this difference in iodine numbers can be tied
up with the phylogeny of the two pines under consideration.

Ivanov (2) has shown that oils of primitive plants had a high
degree of saturation. As evolution of a certain branch of plant
life progressed, unsaturation of the oils increased. According to
Ivanov, probably the iodine numbers of plants of the Carbon-
iferous and Permian had hardly reached 100-120. Progressively
higher iodine numbers appeared later. Although there are many
apparent contradictions to Ivanov’s postulations, there is also
ample experimental evidence to support the theory as a whole.

Ivanov’s theory has found reflection in a recent communica-
tion by McNair (4) briefly abstracted in the American Journal of
Botany. McNair agrees with Ivanov that the “average iodine
numbers increase in value with the increase in evolutionary posi-
tion” of the pines.

In the following table compiled chiefly from Ivanov’s ‘““Vege-
table Oils,” the increase in iodine numbers with evolution of
gymnosperms is shown. The information concerning the geo-
logic age of fossil gymnosperms (represented in the table by
their nearest living equivalents) was supplied by Dr. Ralph W.
Chaney, Professor of Paleontology, University of California,
Berkeley.

Geologic ages of most TeGhats

Species nearly related fossil aR Remarks
gymnosperms

Zamia integrifolia Mesozoic 73

Cycas revoluta Mesozoic 94

Ginkgo biloba Eocene to Miocene 107

Taxodium distichum Eocene to Miocene 107

Pinus monophylla Miocene to Pliocene 108 Adams and Holmes (1)
Pinus sabiniana Pliocene to Pleistocene 112 Author’s data

Pinus radiata Pleistocene 152. Author’s data

From this table it is seen that in gymnosperms the evolu-
tionary development was followed rather closely by increase of
unsaturation of seed oils. The position of Ginkgo in this table is
perhaps questionable, but it should be remembered that the

1938 | SHARSMITH: CLAYTONIA NEVADENSIS 171

Ginkgoales had been developed as an independent branch of
gymnosperms for a very long time. When the iodine numbers
obtained for the two pines under consideration are compared
with the general tendencies of iodine values shown in the table,
the conclusion seems to be in favor of a relatively old age for
P. Jeffreyi.

It appears from the foregoing that in addition to Lemmon’s
“hunch” of a more ancient origin of P. Jeffreyi as compared with
P. ponderosa, we have now both biochemical and distributional
evidence of the relative phylogeny of the two pines.

California Forest and Range Experiment
Station, Berkeley, February, 1938.

LiIrERATURE CITED

1. Apams, Maxwett and Aucust Hotmes. Pine nut oil. Jour. Ind. and Eng.
Chem. 5: 4, 285-287. 1913.

Ivanov, S.L. Vegetable oils. (In Russian) Moscow. 120 pp., illus. 1924.

Lemmon, J. G. Revision of broken cone pines. California State Bd. For-
estry. 3rd Bien. Rpt. 196-201. 1889-1890.

McNair, James B. The phylogeny of Pinus species in relation to glyceride
properties. Am. Jour. Bot. 24: 10, 742. 1937.

Mirov, N. T. Chemical analysis of the oleoresins as a means of distinguish-
ing Jeffrey pine and western yellow pine. Jour. For. 27: 2, 176-187.
1929.

6. Simonsen, JoHN L. and Mapyar Gopat Rav. The constituents of some

Indian essential oils. Indian Forest Records 9 (4): 8. 1922.

SS

pene

ON THE IDENTITY OF CLAYTONIA NEVADENSIS
WATSON

Cari. W. SHARSMITH

Claytonia nevadensis was described by Watson in 1876 from a
specimen collected by J. G. Lemmon in the Sierra Nevada of Cali-
fornia. Since this first description, the species has been mis-
understood repeatedly, mainly because of lack of knowledge or
confusion concerning the nature of the underground parts.
These parts consist of a tangled mass of slender, branching
rhizomes with fibrous adventitious roots, not easily disengaged
from the substratum, especially since the plant usually grows
among rocks in shallow springs and runnels. The type of C.
nevadensis Wats., (Pl. XXIX, fig. 1) upon which Watson’s and
later Rydberg’s (N. Am. Fl. 21: 801. 1982) descriptions were
based, and the type of C. chenopodina Greene (Pl. XXIX, fig. 2)
have been examined. The underground parts of both types,
especially of Greene’s, are meagerly represented so that their
partial or complete misinterpretation by these authors is readily
understandable.

Watson, in the key to the treatment of his type material,
vaguely described the underground system as composed of a
“thickened caudex,’ while in the text he more correctly desig-

172 MADRONO [Vol. 4

nates it as “a rather slender rootstock.” He erroneously de-
scribed the plant as “apparently propagating by runners.” The
term “thickened caudex” gives a misleading picture of the nature
of the underground parts but this part of the description was
perpetuated and accentuated by Rydberg (N. Am. Fl. 21: 297,
301. 1982) by his use of the phrase “‘cespitose rootstock.” In
the type specimen of C. chenopodina Greene, only the upper por-
tion of the slender, branching rhizome remains. Greene, mis-
taking the rhizomes for a root system, described the plant as hay-
ing “‘a fascicle of rather thick and fleshy roots.” The somewhat
fasciculate appearance of the specimen is due to a lateral
rhizome branch which arises in close proximity to a constriction
of the main rhizome. The rhizome branch and the fragment of
main rhizome beyond the constriction are very similar in appear-
ance. They do not appear as if they were originally at all
“thick and fleshy.” The underground parts of this specimen
are typical of those of C. nevadensis, in which the rhizome
branches are occasionally somewhat fasciculate, but there is never
a cluster of thick, fleshy roots as described by Greene.

The new section Chenopodinae in Poellnitz’s treatment of Clay-
tonia (Poellnitz, K. v., Claytonia Gronov. und Montia Mich., in
Fedde, Rep. Spec. Nov. Reg. Veg. 30: 280, 286. 1982) is an ex-
ample of the extent to which an error may be perpetuated. This
section is erected largely on the basis of the “fascicle of several
rather thick and fleshy roots” described by Greene for C. cheno-
podina. “‘Subhastate”’ leaves were also used by Poellnitz as a
sectional character, since Greene had so described the earlier
leaves of C. chenopodina. Poellnitz, however, had not seen
Greene’s type. The leaves present on the specimen are of the
form usual in C. nevadensis.

Claytonia nevadensis Wats. and C. cordifolia Wats., were cited
as synonyms of Montia asarifolia (Bong.) Howell by Gray in the
Synoptical Flora. He indicated C. nevadensis as “‘a reduced
form” of Montia asarifolia. Jepson, in dealing with this problem
(Madrofio 1: 147. 19238), clearly pointed out the differences
which separate Claytonia nevadensis from Montia asarifolia Gray
(sensu lato). As to California specimens, Jepson was dealing
with Claytonia cordifolia Wats., a plant of the northern Rocky
Mountains, Cascades and northern California, probably distinct
from Montia asarifolia (Bong.) Howell. Although separating
Claytonia nevadensis from Montia asarifolia, Jepson noted their
close relationship. His arguments as to the identity of Claytonia
nevadensis as a distinct species are supported by the series of
specimens which have been available to the writer.

A specimen kindly loaned by Dr. Jepson (Mount Leavitt,
Tuolumne County, California, A. L. Grant 420) is the first collec-
tion of C. nevadensis which includes really adequate underground
parts.

1938 ] SHARSMITH: CLAYTONIA NEVADENSIS 173

It is the material upon which Jepson based his concepts in
transferring the species to Montia nevadensis (Wats.) Jepson.
He correctly described the underground system as composed of
“slender, stolon-like rootstocks.” The specimen from Mount
Leavitt clearly shows the rhizomatous condition of the species.
Its rhizomes are slender and abundantly branched, with the
branches occasionally fasciculate along the main rhizome.

The writer was present when the type of Monta alpina East-
wood was collected (Mount Dana, Tuolumne County, California,
in a cold spring on the northwest slope at 11,000—-11,500 feet,
August 5, 1931, H. P. Bracelin 526) and since has made further
observations and collections at this locality over a period of
several seasons. These collections have served as a basis of
comparison with the types mentioned above as well as with the
specimens cited at the end of this paper. The description of
Montia alpina contains an adequate characterization of the
underground parts.

There are two collections on the type sheet of Claytonia
nevadensis, of which the lower, attributed to Lemmon, is the type.
The upper plant (Plumas County, California, August, 1877, Mrs.
Rk. M. Austin) is fragmentary but seems referable to Claytonia
cordifolia Wats. The label under the type specimen reads: “‘Coll.
J. G. Lemmon: Sierra County, &c., 1874.” It was Lemmon’s
custom to put his address on his labels, and he sometimes gave
no clue as to the actual place of collection. Rydberg (N. Am.
Fl. 21: 301. 1982) cites “Sierra Nevada, (side of Mount Dana),
California” as the type locality of the species. He seems to have
had information in addition to what appears on the type label.
Lemmon, in his collecting as state botanist, visited the Mount
Dana region several times. Since Claytonia nevadensis is very
conspicuous in midsummer, growing in dense masses in the shal-
low springs on the northwest plateau of Mount Dana, but is rela-
tively scarce elsewhere on the better known parts of the moun-
tain, it is quite possible that this was the site of Lemmon’s col-

lection and is thus the type locality of Watson’s species as well

as of Montia alpina Eastwood. As further evidence that the
original specimen may not have come from Sierra County, it
may be noted that no other collections of the plant have been
made north of Mount Leavitt in the Sonora Pass region of
Tuolumne County.

The writer would restrict the use of Montia to those forms
with a gamopetalous corolla split on one side and with three
stamens (the M. fontana L. group). Thus considered, the species
under discussion, with its regular petals only slightly united at
the base and its five stamens, falls into Claytonia, if this genus is
viewed in the broad sense as by Poellnitz (Fedde, Rep. Spec.
Nov. Reg. Veg. 30: 280, 286. 1932). The position of the
Species in Claytonia becomes open to question, however, if Ryd-

174 MADRONO [Vol. 4

berg’s construction of the genus is accepted. In the nature of
the underground parts C. nevadensis is closer to Limnia than to
Claytonia as these genera are interpreted by Rydberg. Limnia
is considered by him as having rootstocks and Claytonia as having
corms or thick, fusiform roots. He includes Claytonia asarifolia
and C. cordifolia, both closely related to C. nevadensis in Limnia,
which, according to his conception, has three ovules per capsule
and, in the perennial species, rhizomatous underground parts.
Apparently on the basis of six ovules per capsule he places C.
nevadensis in Claytonia rather than in Limnia. Since C. nevadensis
combines the ovule number of Rydberg’s narrow concept of Clay-
tonia, with the rhizomatous condition of the perennial forms of
Limnia, this species would appear to break down the distinctions
between these two genera as interpreted by him.

In the latest treatment of the Portulacaceae (Pax, F., and
Hoffman, K., Portulacaceae, in Engler and Prantl, Pflanzenf., ed.
2,16C: 257. 1934), there is again an erroneous conception of the
under-ground system of Claytonia nevadensis. It is included with
those species which have a “flaschenférmiger, fleischiger Wur-
zel”’, and placed in section Belia (Steller) Rydb. of Claytoma.
Actually, the nature of the underground system of this species
should place it with Montia as these authors conceive it. Al-
though they place stress upon the underground structures in dif-
ferentiating between Claytonia and Montia, and follow an arrange-
ment different from those of Rydberg and of Poellnitz, these
authors admit the difficulties of the generic problems involved
and the uncertain state of knowledge concerning these groups.
Until generic distinctions in this family are clarified, the writer
prefers to view Claytonia in the broader sense (including Limmnia),
and to delimit Montia as stated above. Presumably differences
in floral structure are of more profound taxonomic significance
than the nature of the underground parts, in which different
forms of storage organs such as rhizomes, corms, or fleshy roots,
may be developed.

CLAYTONIA NEVADENsIS Wats., in Brewer and Watson, Bot.
Calif. 1: 77. 1876. Montia asarifolia Gray Syn. Fl. N. Am. 1:
273. 1897, pro parte non Howell, 1893. Claytonia chenopodina
Greene, Leaflets Bot. Obs. and Crit. 2: 281. 1912. Monta
nevadensis Jepson, Madrofio 1: 147. 1928. Montia californica
Jepson, Man. Fl. Pl. Calif. 349. 1925, as synonym. Montia
alpina Eastwood, Leafl. West. Bot. 1:11. 1982.

Specimens examined: Cauirornia. Sierra Nevada, 1874, J.
G. Lemmon (type of Claytonia nevadensis, Gray Herb.) ; in lava
loam, Mono County, at 10,000 feet, August 4, 1912, Hatton and
Maule (type of Claytonia chenopodina Greene, U. S. Nat. Herb.) ;
foot of Mount Leavitt, altitude 10,000 feet, Tuolumne County,
August 29, 1915, A. L. Grant 420 (Herb. W. L. Jepson) ; alpine |
slopes above Slate Creek, northeast of White Mountain, Mono

175

CLAYTONIA NEVADENSIS

SHARSMITH

1938]

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176 MADRONO [Vol. 4

County, altitude 11,000 feet, in moist, flat, stony streamway, the
roots from beneath stones, September 12, 1929, H. M. Hail
12,755 (Herb. Univ. Calif.) ; in a cold spring on the northwest
slope of Mount Dana, Tuolumne County, at 11,000—-11,500 feet,
August 5, 1931, Mrs. H. P. Bracelin 526 (type of Montia alpina
Eastwood, Herb. Calif. Acad. Sci., No. 189,088) ; in dense masses
in shallow springs, northwest plateau of Mount Dana, altitude
11,500 feet, Tuolumne County, August 9, 1933, C. W. Sharsmith —
342 (Herb. Univ. Calif.) ; same locality, September 10, 1934, C.
W. Sharsmith 2019 (Herb. Univ. Calif.) ; same locality, September
18, 1936, C. W. Sharsmith 2324 (Herb. Univ. Calif.) ; Tioga Crest,
east of Saddlebag Lake, altitude 11,400 feet, Mono County, July
31, 19386, H. L. Mason 11,462 (Herb. Univ. Calif.).

University of California,
Berkeley, May 12, 1937.

PICEA BREWERIANA IN SHASTA COUNTY

Puitie G. Happock

Rumors of the occurrence of the weeping spruce (Picea
Breweriana Wats.) in the Mount Shasta region are verified by
the finding in May, 1936, of a small grove of the species on Castle
Crags in the northwestern corner of Shasta County. The group
occurs at an elevation of approximately six thousand feet and is
near the quarter-section corner between sections 4 and 5, Town-
ship 38 N., Range 4 W. of the Mount Diablo Meridian. All of
the trees seen are located in the saddle or in the gulch to the
north between the main higher group of crags and the dome-
like rock termed Castle or Battle Rock, which stands out a little
to the east of the others. It is estimated that there are about
twenty individuals, which would average fifty feet in height and
up to two feet in diameter, breast high. They are associated
with Abies magnifica, Pinus Jeffreyi, P. Lambertiana, Arctostaphylos
nevadensis, A. patula, Castanopsis sempervirens and Ledum glandu-
losum.

This spruce is one of the most restricted in range of any
species of Picea. It occurs in greater abundance to the northwest,
in southern Oregon, and to the southwest, in Trinity County, Cali-
fornia. The Castle Crag locality is perhaps the most easterly
station. Unfortunately, the boundary of Castle Crag State
Park is so situated that this grove is narrowly excluded from the
preserve.

Berkeley, California,
July 22, 1937.

1938] BUTTERFIELD: ACACIA IN CALIFORNIA 177

THE INTRODUCTION OF ACACIAS INTO CALIFORNIA

H. M. BurtrerrFrie_p

A visitor to almost any city in the milder districts of Califor-
nia, from San Diego County north to Sonoma County and inland,
is impressed with the fact that two of the most common orna-
mental trees are exotics. The acacia and the eucalyptus were
brought in so many years ago that they appear quite as much at
home as most native trees. Previous to 1849 there were no Aus-
tralian acacias in California. It may be true that Acacia Far-
nesiana was grown about the Spanish misisons at San Diego and
Santa Barbara’ at an earlier date, yet this species is often con-
sidered American. The Spanish explorers apparently found it
at an early period and ultimately brought it to California by way
of Mexico. Paxton in the Botanical Dictionary listed this species as
having been introduced from San Domingo in 1656. This fact
tends to show that early writers associated the first plants of
Acacia Farnesiana with America, even though the species is now
found in many countries.

When gold was discovered in California, people were at-
tracted from all parts of the world. Australians probably
brought seeds of the acacia and supplied them to early growers.
California nurserymen were soon importing seeds directly from
Australia. Many facts regarding these early importations are
now available.

Some of the first nurserymen in California had been trained
in Europe and were familiar with the acacias already introduced
into England. Other nurserymen had English reference books
which listed many species. Perhaps most readers of today
would be surprised to find a long list of acacias in Paxton’s
Botanical Dictionary, published in London in 1849. When Wil-
liam C. Walker, one of the earliest growers of acacias in Califor-
nia, was making up his catalog for 1858-59 he wrote regarding
certain species of Australian trees “not in Paxton’s” so we know
that these early California nurserymen did use references to
check on the acacias which were being imported. Ferdinand
von Mueller also began to write of his discoveries in Australia
before 1860 so early California nurserymen were able to read
about many species of Acacia.

Botanical publications furnish dates when various species of
Acacia were introduced into England. For example, we find that
A. verticillata was imported into England as early as 1780, A.
pubescens in 1790, A. longifolia in 1792, A. armata in 1808, A. de-
currens var. mollis in 1810, A. melanoxylon in 1819, A. cultriformis
and A. decurrens var. dealbata in 1820. We would naturally ex-

1 Shinn, Chas. H. An economic study of acacias. U.S. Dept. Agric. Bull.
9: 1-38. 1913.

178 MADRONO [Vol. 4

pect the first California nurserymen to include these species in
their first plantings. That was exactly what happened.

To look back over a period of about eighty-five years and say
that certain species of acacias were the first to be planted in Cali-
fornia may be difficult, yet it is possible to offer some interesting
facts. Colonel J. L. L. F. Warren, the editor of The California
Farmer, a paper started in 1853, was also a Sacramento nursery-
man. He visited various nurseries and gardens in central Cali-
fornia and reported his observations. After a visit to the garden
of Mrs. C. V. Gillespie in San Francisco, he reported that the
acacias grown by this lady were the first in California. During
the early fifties Colonel Warren’s nursery was at 15 J Street,
Sacramento. His catalog was issued here under the name War-
ren and Sons Garden and Nurseries. A copy of this catalog for
1855, located in the Bancroft Library of the University of Cali-
fornia, lists “Acacia armata,” “Acacia dealbata,” “Acacia florabunda,”
and “Acacia longifolia.” The spelling of Acacia longifolia var.
floribunda does not agree with modern usage and Acacia dealbata
is now referred to Acacia decurrens var. dealbata, but all of the
above acacias are easily recognized today. These four species,
listed within perhaps two or three years of their introduction,
are probably the first Australian acacias in California. Acacia
seeds were being advertised for sale in 1854 by Colonel Warren,
according to Charles H. Shinn*. Colonel Warren is also reported
to have sold plants of Acacia decurrens and A. melanozylon. His
catalog of 1853-54 does not include A. melanoxylon which was,
however, listed before 1860.

Acacias sold in early days were soon widely distributed. Ac-
cording to the The California Farmer, E. L. Beard had two trees
at Mission San Jose in 1855; Dr. Cobb of San Jose had an acacia
tree in 1855 that was already 15 feet high and had a spread of
25 feet; Captain Walsh of Benicia and Captain C. M. Webber,
founder of Stockton, each had an acacia tree in 1855. The
Stockton tree, the editor stated, was only a year old. This would
suggest that the others were older.

Reminiscences of early residents are useful in determining
dates of introduction, yet memory is sometimes faulty. Lists in
old nursery catalogs and advertisements in old papers furnish evi-
dence that is largely beyond dispute. In addition to the catalog
of Warren and Sons for 1858, there is a catalog of the ‘Shell
mound Nurseries and Fruit Gardens” for the year 1856 in which
the Australian acacia is listed at $1.00 for plants 12 to 18 inches
high. R.W. Washburn was owner and a Mr. Sanford manager
of the nursery which was located near Brooklyn, now a part of
Oakland, California.

Two brothers, William and James O’Donnell, had a nursery
on Mission Street, San Francisco, in the early fifties named “The
United States Nursery.” According to The California Farmer
they had in 1856 “two acacias of rare beauty and size’ which

1938] BUTTERFIELD: ACACIA IN CALIFORNIA 179
they sold at $50 each. In the same year William O’Donnell
moved to San Jose, where he established the “Mountain View
Nursery” at 532 William Street between Ninth and Eleventh
streets, and there continued to grow the acacia. By 1858, ac-
cording to the report of the California State Agricultural Society
for that year he was growing five thousand acacias.

William C. Walker operated his “Golden Gate Nursery” at
Fourth and Folsom streets, San Francisco, until after 1865. He
began advertising the acacia as early as 1857 when he reported
ten kinds. Until the master copy? of his 1858 catalog was lo-
cated the species which he offered for sale were unknown. Mr.
Walker also listed the acacia seeds brought in late in 1859. But
let us go over the species offered for sale in 1858, giving them
in the exact order found in the old catalog and with the origina]
spelling. Synonyms recognized today are given in brackets.

Acacia lophantha [ Albizzia Acacia julibrissin [ Albizzia

lophantha | julibrissin |
2 speciosa [ Albizzia = dodoenifolia [ A.
Lebbek | dodonaefolia]
“ leptophylla [ A. “ cornigera
. Farnesiana | ‘“ suaveolens

T4

armata

dealbata [ A. decurrens
var. dealbata |

latifolia

verticillata

salicifolia [ ?]

capensis [ A. horrida |

longifolia

strombulifera [ Prosopis
strombulifera |

falcifolia [ A.

procumbens |
Farnesiana
linearis or longissima

trinerva [ A. trineura or
A. trinervata ?]
rugosa [ A. rugata or
A. concinna ?]
melanoxylon _ [ Walker’s
synonym of A. latifolia
probably incorrect |
vestita
barbon (Poinciana Gil-
lesii) [this is Walker’s
listing |

Silver Wattle [ A. decurrens var.

dealbata |

Acacia heterophylla

The botanical names given by Mr. Walker may seem a little
out of date at this time and yet up to 1900 or later the species of
Albizzia were still being listed under the genus Acacia. Any stu-
dent of botany who cares to look up the names offered by Mr.
Walker will realize that early nurserymen were very well in-
formed for their day, in fact few references of today will include
all of the acacias listed in Mr. Walker’s catalog of 1858 to 1861.

In 1857 Mr. Walker advertised ten kinds of acacias and five
hundred plants. By 1858 the number of species had been more
than doubled. By 1861 Mr. Walker was advertising seventy spe-

2Copy of Walker’s catalog in possession of Miss Alice Eastwood, Cali-
fornia Academy of Sciences, Golden Gate Park, San Francisco.

180 MADRONO [Vol. 4

cies and varieties. No California nurseryman since his time has
listed so many Australian acacias. New species have been intro-
duced from time to time but Mr. Walker was one of the greatest
specialists in growing acacias that California has ever had. Mr.
Walker continued to issue nursery catalogs until 1865 or later
and fortunately copies of his 1860, 1861 and 1865 catalogs have
been preserved from the estate of Captain C. M. Webber. A list
therefore of the seventy kinds of acacias advertised can now be
presented. To make checking easier, the species listed by Mr.
Walker are here arranged alphabetically with synonyms in
brackets. Some species and varieties listed are still of doubtful
identity ; evidently Mr. Walker listed the same species under dif-
ferent names, not realizing at the time that they were synony-
mous. In reality there were not seventy distinct species, yet every
reader will marvel at the long list of acacias, several of which are
still rare or unusual. The following species were listed by Mr.
Walker in 1860 and 1861:

Acacia argyrolofera [A. brachy- Acacia floribunda latifolia ?
botrya var. argyro- - glandulosa [Prosopis

phylla ?] glandulosa]
armata : glauca pendula [A. pen-
“_ bienana [ A. bynoeana ?] dula var. glaucescens ? |

bispinata [ A. bispinosa ] grandis [ A. pulchella var.
caesia grandis |
capensis [ A. horrida| Guilfoyliana [ possibly va-

7 celastrifolia [ A. myrtifolia riety named after M.
var. celastrifolia | Guilfoyle |

¥ chordaphylla [ A. rigens | pe heteroclada [ A. hetero-

4 cornigera - clita ?]

i cultriformis : heterophylla

a cuspidata [ A. diffusa var. . hybrida [ A. armata |
cuspidata | Pe Hugelii [ A. Huegelii]

4 dealbata [ A. decurrens i impressa
var. dealbata | a ixiophylla

i decipiens a julibrissin [ Albizzia juli-

; decurrens [A. decurrens brissin |
var. normalis | ia latifolia

ce dodoenifolia [ A. dodonae- - La Trobei [A. acinacea
folia | var. Latrobei |

z Douglasii [listed by Pax- 3 leptophylla [ A. Farnesi-
ton, 1868 | ana |

a exudens [ A. exsudens, A. “ leucophylla [ Leucaena
verniciflua | glauca |

falcifolia [ A. procumbens | i linearis

i Farnesiana : linifolia

i floribunda [A. longifolia ‘ longiflora floribunda [A.
var. floribunda | longifolia var. flori-

floribunda pendula ? bunda |

1938 ] BUTTERFIELD: ACACIA IN CALIFORNIA 181

Acacia longiflora magnifica [pos- Acacia pudalosa [ A. paludosa, A.

sibly a variety of A. pennata |
longifolia | iy pulchella
fe longifolia a pulchella spinosa [a vari-
‘“ Jongissima [ A. linearis | elie
. lophantha [ Albizzia lo- ‘ pyracantha [A. pycnan-
phantha | tha ?]
“ melanoxylon “rotundifolia [A. obliqua]
‘““ mimeroides [ ?] _ rugosa | A. rugata, A. con-
: molissima [A. decurrens cinna ? |
var. mollis | i salicifolia [ A. salicina ? |
“montana i. sophora | A. longifolia var.
‘“ monospermum [| ? | sophorae |
“mucronata [A. longifolia a speciosa [Albizzia Leb-
var. mucronata | bek ]
“ myrtifolia spectabilis
;° myrtifolia elegans [ vari- i‘ strombulifera [ Prosopis
ety of A. myrtifolia ?] strombulifera |
i; nigricans suaveolens
: neiomanii [ ? | Bs trinerva [ A. trinervata, A.
o paradoxa [ A. armata | trineura, A. trinervia ?]
yi pendula st verticillata
% pubescens ‘L vestita

A word concerning the prices charged by Mr. Walker may be
of interest. In 1858 the following prices were asked: Acacia
linearis and Albizzia Julibrissin, $5; Acacia decurrens var. dealbata,
$3-$5; A. longifolia, $2.50; A. verticillata and A. dodonaefolia, not
priced. In 1860 prices had dropped materially and were quoted
as follows: Acacia armata, $1; A. cultriformis, $1.50; A. decurrens
var. dealbata, $2.50; Albizzia Julibrissin, $2.50; Acacia verticillata,
$1.50. Others were not priced, indicating that they may have
been scarce.

Seeds of acacia were received by Mr. Walker from Mr. M.
Guilfoyle who had established his “Exotic Nursery” at Double
Bay, Sydney, about 1852. (His son, W. R. Guilfoyle*®, did much
to establish the Melbourne Botanical Gardens.) It is possible
that Mr. Walker had heard of M. Guilfoyle’s nursery through Dr.
Herman Behr who had spent several years in Australia and who
was active in the Academy of Sciences, San Francisco, as early as
1854. The following kinds of acacia seed were received by Mr.
Walker from M. Guilfoyle in 1859 and were later made available
for sale: Acacia cultriformis, A. spectabilis, A. decurrens var. mollis,
A. verticillata, and A. diffusa var. cuspidata. Mr. Walker issued a
catalog as late as 1865; his name disappeared from the San Fran-
cisco directory about 1867.

The Suscol Nurseries established by Simpson Thompson in
1853 about six miles south of Napa where, some time before, land

3 Information received by author from M. Guilfoyle’s daughter, 1935.

182 MADRONO [Vol. 4
had been secured from General Vallejo, are also of interest in
connection with the acacia. This nursery sold acacias in 1858,
but judging from the catalog of 1861, did not continue to handle
them.

About 1860 Stephen Nolan of Oakland established his Belle-
view Nursery and there started to raise the acacia. Mr. Nolan
was born on the Island of Jersey in 1818, and since he was ap-
prenticed in England, he was doubtless familiar with the Aus-
tralian acacia when he came to California. Before starting his
nursery, Mr. Nolan had worked on the Potter estate, and the fact
that he later listed a plant, which he considered a hybrid, as
Acacia Potteri might indicate that he had worked with acacias
while there. In addition to growing acacias in his nursery, Mr.
Nolan also introduced eucalyptus* about 1860, and in a short time
he became one of the most important growers of Australian
plants that California has ever had. While we do not know all
that Mr. Nolan handled in the sixties, we do have his catalog for
1871°. He listed thirty-four species of acacias in that year.
Prices ranged from twenty-five cents to seventy-five cents each
for most kinds. The following list of species grown by him may
be of interest:

Acacia armata [also listed by Acacia leucantha [A. leucacan-

Nolan as A. undulata | tha ?]

a calamifolia i linearis

4 celastrifolia [ A. myrtifolia a longifolia
var. celastrifolia | ie melanoxylon

chordophylla [ A. rigens ]

cultriformis

dealbata [| A. decurrens
var. dealbata |

floribunda [A. longifolia
var. floribunda |

homalophylla

imbricata [ A. lineata]

implexa

latifolia

leiophylla [ A. saligna |

mollisima [A. decurrens
var. mollis |

prominens [A. linifolia
var. prominens |

pycnantha

reclinata [ A. leprosa]

retinodes

Sophorae [A. longifolia
var. Sophorae |

trinervata

verrucosa [ A. venulosa ?]

In attempting to give synonyms for old names of Acacia there
is always some possibility of error. Nurserymen, printers, and
others have always had trouble in copying correctly and for cer-
tain species several names have been in use. Acacia floribunda of
the gardens in early days may have been A. retinodes. A. flori-
bunda is often referred to A. longifolia var. floribunda. Today we

4 Butterfield, H. M. Introduction of Eucalyptus into California. Madrofio

3: 149-154. 1935.
5 Copy of the catalog in possession of Mrs. W. Ee Snyder of Berkeley,
daughter of Stephen Nolan.

1938] BUTTERFIELD: ACACIA IN CALIFORNIA 183

have both A. retinoides var. floribunda and A. longifolia var. flori-
bunda so we can only guess at what early nurserymen meant.
Acacia Reimeri, possibly named for E. L. Reimers of San Fran-
cisco, was mentioned in early literature. This acacia may have
been a seedling that differed from the parent plant, or the origi-
nal label may have been lost and a new name substituted. Con-
siderable variation exists in certain species, and hybrids have
been found from time to time, so it would not be strange if these
early nurserymen had difficulty in properly classifying certain
seedlings; an easy solution was to give a new name.

During the sixties certain additional species of Acacia were
doubtless imported but little is known about this period. Julius
Ferrer’ of San Francisco, according to his report in the Pacific
Rural Press at a later date, was growing at his nursery in 1862
the following species: A. cyanopyhlla, A. cuneata, A. decurrens vay.
dealbata, A. linearis, A. longifolia, A. linata, A. melanoaylon, A.
decurrens var. mollis, A. pendula, and A. Riceana.

Most of the early acacias in California were started from im-
ported seed but some plants also were brought in. As early as
1859 Mr. Walker brought in to San Francisco Acacia pubescens in
a Wardian case. George Gordon, nurseryman of Menlo Park,
imported some trees of Acacia latifolia about 1869. Unfortunately
these plants were infested with the cottony cushion or white scale
(Icerya purchasi)®. Oranges brought in from Australia at the
same time also became infested and, when shipped to southern
California, spread this serious insect pest to citrus orchards. It
gradually spread throughout the state and by 1877 was attacking
acacias in Marin County. Perhaps this experience aided in the
development of mineral oil sprays, and it did lead to the intro-
duction from Australia of the Vedalia (Rodolia cardinalis), a small
red ladybird beetle which preys on this scale.

The change that took place in California gardens from 1860
to 1880 was remarkable. Dr. Herman H. Behr, a friend of Ferdi-
nand von Mueller, who came to California in 1851, became asso-
ciated with the California Academy of Sciences in San Francisco
and was instrumental in introducing the various Australian plants
into California. In 1870 W. H. Hall started acacia plantings in
Golden Gate Park, setting out twelve hundred trees, representing
ten species’. While Dr. Behr did not do any of the planting, his
friendship with von Mueller probably had something to do with
the introductions of the acacias. In 1880 Dr. Behr was led to
say, “The vegetation of the peninsula [ referring to the peninsula
from San Francisco south through San Mateo and Menlo Park]
is at present more Australian than Californian.” In later years
as many as fifty thousand acacia trees were set out in Golden
Gate Park in a single year, partly to help bind the sand and

6 Essig, E. O. A history of entomology. p. 119. Macmillan Co., 1931.
* Pacific Rural Press, San Francisco, February 21, 1880.

184 MADRONO [Vol. 4

partly as ornamentals. Acacia longifolia was used extensively for
this purpose. In time perhaps half a million acacia trees, repre-
senting sixty species, were planted in Golden Gate Park. Mr.
John McLaren should be honored for his part in this enterprise.
Some of these Acacia species® are listed below: A. acinacea, A.
acuminata, A. adunca (crassiuscula), A. aestivalis, A. alata, A. armata,
A. aspera, A. Baileyana, A. Betchei, A. Cavenia, A. confusa, A. cultri-
formis, A. Cyclops, A. decurrens var. dealbata, A. decurrens vay.
mollis, A. dentifera, A. Dietrichiana, A. dodonaefolia, A. elata, A.
Farnesiana, A. fimbriata (form of A. linifolia var. prominens), A.
glaucescens, A. hastulata, A. implexa, A. iteaphylla, A. juncifolia,
A. Koa, A. leprosa, A. lineata, A. longifolia, A. longifolia var. flori-
bunda, A. longifolia var. latifolia, A. longifolia var. Sophorae, A.
melanoxylon, A. microbotrya, A. obliqua, A. oxycedrus, A. pendula,
A. pentadenia, A. podalyriaefolia, A. pravissima, A. prominens (A.
linifolia var. prominens), A. pruinosa, A. pulchella var. hispidissima,
A. pycnantha, A. retinodes, A. retinodes var. floribunda, A. retinodes
var. gracillima, A. Riceana, A. salicina var. Wayae, A. saligna, A.
stenoptera, A. tenuifolia, A. verticillata.

Several nurserymen sold acacias in San Francisco between
1870 and 1880. E. L. Reimers exhibited several of his acacias at
the Bay District Horticultural Fair held in San Francisco in 1871,
among them Acacia conspicua (A. vestita), A. cordata (trade name
not recognized), A. alata, A. pulchella, and A. rubra (A. rubida ?)?°.
He had also many of the old favorites.

The Pacific Nursery of San Francisco, operated by F. Liide-
mann, included the following species of Acacia in a catalog for
1874—75'°: A. albicans [ Pithecolobium albicans], A. conspicua [ A.
vestita|, A. farinosa, A. fragrans [| Albizzia fragrans|, A. La Trobay
[ A. acinacea, A. Latrobei].

With the establishment of the College of Agriculture at the
University of California in 1874 a new influence was exerted on
horticulture. Landscape men of the old school actually objected
to the large number of Australian trees set out on the new
campus. Dr. Eugene Hilgard arrived in 1875 to serve as Direc-
tor, and E. J. Wickson joined him later. Both took an active
interest in the distribution of acacias to people throughout Cali-
fornia. All through the earlier days of the University standard
species such as A. melanozylon and A. decurrens or its varieties
received much attention. <A total of five thousand individuals and
25 species had been planted on the campus by 1875. Acacia
seeds of new species, such as A. pulchella var. grandis, were re-
ceived as early as 1879. In time the institution attracted the
attention of such men as Baron von Mueller. We find in the re-
port of the Experiment Station for 1886 that this noted botanist

8 List supplied by Miss Katherine D. Jones, College of Agriculture, Uni-
versity of California.

9 For detailed list see “The California Horticulturist,” 1871.

10 Catalog in the Bancroft Library, University of California.

1938 ] BUTTERFIELD: ACACIA IN CALIFORNIA 185

sent seeds of A. sentis and A. spectabilis. It was also in 1886
that seeds of A. arabica were received. Dr. Herman Behr and
others doubtless assisted in forwarding acacia seeds for further
increases. Two years later an inventory of the trees on the
campus showed well over twenty species of acacias. Most of
those grown were early introductions, but a few such as A.
Cavenia, A. Greggit, A. imbricata (A. lineata), and A. Riceana were
not so widely known. The planting of new species of acacias on
the campus at Berkeley continued until well after 1900. Acacia
elata was planted in 1901.** Specimens of Acacia cultriformis and
several other species planted at about this time are still living.

In 1885 forestry experiment stations were established at
Chico and Santa Monica under the State Board of Forestry.
Some acacias were set out before the University of California
took over the stations in 1898. At that time A. retinodes, A. cy-
anophylla, and A. latifolia were doing well but probably many
trees had succumbed because of poor care. Planting continued
after the University took over the work, especially during the
years 1900 to 1910. A survey of the trees at the Santa Monica
Forestry Experiment Station in 1917 by W. Metcalf, now exten-
sion forester in the College of Agriculture, University of Cali-
fornia, showed the following species: A. arabica, A. Baileyana, A.
binervata, A. cultriformis, A. cyanophylla, A. Cyclops, A. decurrens,
A. decurrens var. dealbata, A. decurrens var. mollis, A. Dietrichiana,
A. elata, A. Jonesii, A. leprosa, A. longifolia, A. longifolia var. flori-
bunda, A. Maideni, A. melanoaylon, A. neriifolia, A. pruinosa, A. pyc-
nantha, A. saligna, A. verniciflua (listed as A. virgata).

Conditions at the Chico Forestry Experiment Station were not
so favorable for acacias and yet attempts were made to grow
many species. Some of those growing in 1893 in the nursery”
are: A. arabica, A. argyrophylla (A. brachybotrya), A. Bartheriana
(A. Berteriana), A. capensis, A. cyanophylla, A. dodonaefolia, A.
glauca (A. glaucescens), A. glomerata (A. glomerosa ?), A. holo-
cericena (A. holocericea ?), A. leucocephala (Leucaena glauca), A.
lophantha (Albizzia lophantha), A. lunata, A. melanozylon, A. mono-
phylla (?), A. ornithoflora (A. armata ?), A. ovalifolia (?), A. pere-
grina (Piptadenia peregrina ?), A. pycnantha, A. spectabilis var.
excelsa, A. trinervis (A. trineura or A. trinervata ?).

For several years after 1885 nurserymen showed very little
interest in new species of acacias. Here and there a nurseryman
would include some new species, but no large number was added.
John Rock, who came to San Jose in 1865 and helped start the
California Nursery at Niles in 1885, had doubtless seen California
horticulturists wax enthusiastic about Australian acacias, yet he
always showed only a moderate interest by listing a few popular

species. In 1888 Mr. Rock included A. falcata in his catalog.

11 Jones, Katherine D. Acacias in California. National Horticultural
Magazine. January, 1933.
12 California Agricultural Experiment Station Report. 1893.

186 | MADRONO [Vol. 4

Many years later, in 1915, his successors were still adding a few
new species such as A. Baileyana, A. juniperina, A. neriifolia, and A.
podalyriaefolia, but the additions were made cautiously and in
accordance with demand. The handling of new species of orna-
mentals is not always profitable. Introduction of other new spe-
cies had to await the interest of enthusiastic plant importers such
as Dr. F. Franceschi (Dr. F. Fenzi) and E. O. Orpet of Santa
Barbara, or of horticulturists connected with some institution not
dependent on the financial outcome, such as Golden Gate Park >
or the University of California. |

The history of the introduction of acacias into California is
by no means limited to central California, even though the first
acacias were grown near San Francisco and Oakland. Several
nurserymen and private growers in Santa Barbara, Los Angeles,
and San Diego counties have imported new species. The follow-
ing listed by Dr. Franceschi and P. Reidell in their catalog for
1908 and later were of special interest: A. accola, A. Baileyana, A.
elongata, var. angustifolia, A. extensa, A. leptoclada, A. obliqua, A.
pendula, A. penninervis, A. podalyriaefolia, A. pravissima, A. subulata.
Other plants listed with these in the 1908 catalog were Pitheco-
lobium flexicaulis, Acacia corymbosa and A. notabile (A. notabilis ?).

Mr. E. O. Orpet of Santa Barbara has grown some interesting
species of Acacia in recent years, including A. Bakeri, A. obtusata,
and A. mucronata. Acacia pulcherrima reported growing at Mr.
Orpet’s nursery is probably Stryphnodendron floribundum. :

Still another important center of introduction in southern
California is to be found about San Diego where Miss Kate Ses-
sions, T. Wayland Vaughn and others have shown an active inter-
est in novelties. Mr. Vaughn has been associated with Scripps’
Institute of Oceanography where such species of Acacia as A. car-
diophylla, A. decurrens var. Leichardtu, A. falcata, A. flexifolia, A.
hakeoides, A. leptoclada, A. linifolia var. prominens, A. nonuttiana
(?), and A. viscidula have been grown. Other Acacia species
reported about San Diego include A. aneura, A. Bancrofti, A.
Betchei, A. decora, A. decurrens var. pauciglandulosa, A. penninervis,
and A. polybotrya var. foliolosa. The San Diego climate should
prove to be very favorable for some of these less common species.

The United States Department of Agriculture, through the
Office of Foreign Seed and Plant Introduction, has imported cer-
tain species of Acacia. Several of these are native to Africa and
are of questionable ornamental value. Others listed have been
grown previously, but the reintroduction of Acacia species has
taken place many times and we can expect many will be reintro-
duced in the future. Perhaps only a few of the three hundred
Australian species are really adapted to California and only a
few will live for any long period.

Inquiry is often made as to the location of old acacia trees in
California. Most acacias are shallow-rooted and interfere with
the growth of other plants near by; they also clog sewers at

1938 ] MASON: HARRY STANLEY YATES 187

times and break sidewalks. For such reasons the trees are often
removed while still young. Low winter temperatures which
occur occasionally have wiped out many acacias, the freeze of
1913 having killed most of those at the Chico Forestry Experi-
ment Station, and the freeze of 1933 having killed all of the
acacias on the State Capitol grounds at Sacramento. Very old
acacias have been removed in cities as business enterprises grew
and needed space, and even under favorable conditions some of
the species introduced would not have survived very long in Cali-
fornia. These several factors help to account for the scarcity of
old acacias in the state.

A few old trees of Acacia melanozylon planted in 1879 are still
growing along Berkeley streets. One of these trees now
measures 2.9 feet in diameter breast high and is 70 feet tall. Old
trees of Acacia verticillata are also found in Berkeley. Probably
old acacia trees are still alive in other parts of California.
Readers are invited to report such specimens, giving the age and
any interesting facts associated with their introduction. Readers
can assist also in completing this story of the Australian Acacia
in California by reporting rare species not included in this record.

College of Agriculture,
University of California,
November, 1936.

HARRY STANLEY YATES

Harry Stanley Yates, the only child of Francis Frederick Yates
of Kidderminster, England, and Jennie Gibson Yates of Wor-
cester, Massachusetts, was born in Marshall, Minnesota, on Oc-
tober 2, 1888, and died in Berkeley, California, January 17, 1938.
The family came to California in 1890 and settled on an orange
ranch near Pomona. In 1912 he was married to Alice B. Weber,
a botanist and a fellow member of the University of California
class of 1912. Dr. Yates is survived by his wife, his daughter,
Elizabeth Yates Biernoff, and his son, Francis Gordon Yates.

After graduating from Pomona High School in 1908, he at-
_ tended the University of California at Berkeley, graduating in
1912. The degrees Master of Arts and Doctor of Philosophy
were conferred upon him by the University of California in 1914
and 1915. He was a life member of the Malayan Branch, Royal
Asiatic Society, and of the National Geographic Society, a charter
member of the California Botanical Society, and a member of the
University of California chapter of Sigma Xi.

Upon receiving his doctor’s degree he accepted a position as
mycologist for the Bureau of Science, Manila, Philippine Islands.
After spending five years in this position he became research
botanist for the United States Rubber Company on its plantation
at Kisaran, Asahan, Dutch East Indies, East Coast Sumatra. In
1929 he returned to the United States with his family, and in

188 MADRONO [Vol. 4

1932 became associated with the California Forest Experiment
Station as botanist for the Vegetation Type Survey of California
and western Nevada. In this work he was responsible for train-
ing the field men in plant identification, for checking the field
determinations and for the organization and development of an
herbarium for the permanent preservation of the specimens upon
which the maps are based. At the time of his death this her-
barium contained a collection of more than twenty-three thousand
plants. |

From his student days, Dr. Yates was an enthusiastic collector,
some eight thousand specimens having been donated to the Uni-
versity of California Herbarium as a result of his labors. His
earlier collections were largely fungi, while his later ones were
more general. In 1915 he made a valuable collection of the
plants of the Trinity National Forest, California. In the Philip-
pines he continued his interest in general collecting, but his best
known collections are those made in Sumatra, a set of six thou-
sand flowering plants and one thousand fungi of this series being
deposited in the University of California Herbarium. In recent
years, he became especially interested in California grasses, and
his extensive collections are a distinct contribution to our know]l-
edge of the grasses of this state. The large collections of the
genus Arctostaphylos that passed through his hands in connection
with his duties in the Vegetation Type Map Survey gave him a
very complete knowledge of this genus. Although the work is
unpublished, it was organized for use by the field crews and
proved to be a very workable treatment of a difficult group.—
Hersert L. Mason.

List oF PusBLISHED WRITINGS

1915. Stock Poisoning Plants of California (with H. M. Hall). Calif. Agric.
Exp. Stat. Bul. No. 249: 219-247.

1916. The Comparative Histology of Certain Californian Boletaceae. Univ.
Calif. Publ. Bot. 6: 221-274, pls. 21-25.

1917. Copra and Coconut Oil (with H. C. Brill and H. O. Parker). Philippine
Jour. Sci. 12: 55-86.

Fungi collected by E. D. Merrill in southern China. Philippine Jour. Sci.
12: 313-316.

The revegetation of Volcano Island, Luzon, Philippine Islands, since the
eruption of Taal Volcano in 1911 (with W. H. Brown and E. D.
Merrill). Philippine Jour. Sci. 12: 177-248, pls. 4-16.

The rate of growth of some trees on the Gedeh, Java (with W. H.
Brown). Philippine Jour. Sci. 12: 305-310.

Some recently collected Philippine fungi. Philippine Jour. Sci. 12:
361-380.

1918. Fungi from British North Borneo. Philippine Jour. Sci. 13: 233-240.

Some recently collected Philippine fungi, II. Philippine Jour. Sci. 13:
361-384.

1919. The growth of Hevea brasiliensis in the Philippine Islands. Philippine
Jour. Sci. 14: 501-523.

Pink Disease of Citrus (with L. H. Atherton). Philippine Jour. Sci. 14:

657-671, pls. 1-7.

1938 | FOSBERG: ERIOGONUM ABERTIANUM 189

ERIOGONUM ABERTIANUM AND ITS VARIETIES

F. Raymonp FossBere

Dr. John Torrey described Eriogonum Abertianum in Major
Emory’s “Notes of a Military Reconnoissance” (p. 151, 1848).
The plant is an annual, ordinarily dichotomously or trichoto-
mously branched near or above the base, canescently tomentose
to villous, with campanulate involucres bearing many flowers;
the perianth parts are in two series, the outer three expanded,
more or less orbicular, membranous-scarious, covering the narrow
inner ones. This species is closely related to E. pharnaceoides
Torr., differing in the pubescence, the leaf shape, and in the more
expanded outer perianth segments which are thinner and more
scarious. It is also related to E. ovalifolium Nutt., from which it
differs in being an annual, in the pubescence and shape of its
leaves, and in the open rather than condensed inflorescence.

A study of material from the Mesilla Valley, New Mexico,
indicated that there were two different entities which keyed, in
the Flora of New Mexico by Wooton and Standley, to Eriogonum
Abertianum. In the synonymy was given the name Lriogonum
cyclosepalum Greene (Muhlenbergia 6: 1. 1910) the description
of which seemed to fit one of the Mesilla Valley plants. The
remainder of the material agreed better with E. pinetorum as de-
scribed by Greene in the same paper than with E. Abertianum
Torr. as interpreted by him.

In an effort to settle the problem, Eriogonum Abertianum and
related species were studied in the herbaria of Pomona College,
California Academy of Sciences, Los Angeles Museum, and the
University of California, Berkeley. The material examined in-
cluded isotypes and cotypes of E. pinetorum Greene, E. cyclosepa-
lum Greene, EL. arizonicum Gandoger, E. Abertianum var. neome.si-
canum Gandoger, and E. Abertianum var. ruberrimum Gandoger.
Dr. Gleason had photographed for me at the New York Botani-
cal Garden the type of E. Abertianum in the Torrey Herbarium.

As shown by these photographs, Dr. Torrey had at hand only
one sheet which bears material collected by Abert. This sheet
contains two specimens, one a small fragment of the top of a
plant, labelled “July 17, Lt. Abert,” and another marked “Oct.
14th, 1846, Emory.” <A careful study of Emory’s Report shows
that on July 17 Lieutenant Abert was near the junction of the
Pawnee River with the Arkansas River, in Pawnee County, Kan-
sas, considerably out of the present range of the species, and on
October 14, Emory was along the Rio Grande, apparently in
Sierra County, New Mexico. Both of these plants seem to be
Eriogonum Abertianum var. neomezicanum Gandoger. The other
sheet in Torrey’s herbarium bears four specimens. The plant on
the left of the sheet, no. 1, collected in August by Dr. Bigelow
from “Near San Diego (Vall. of R. Grande)” is E. Abertianum

190 MADRONO [Vol. 4

var. bracteatum. The remaining three specimens should all be
referred to E. Abertianum var. cyclosepalum. They were collected
in Mexico by Dr. Parry: no. 2 at Chihuahua; nos. 3 and 4 from
Janos [Janas?], northern Chihuahua. All of these plants were
named simply “Eriogonum Abertianum” by Torrey.

The groups of plants in this complex are best treated as varie-
ties of a single species, Eriogonum Abertianum Torr., because of
the insignificance of the characters in which they differ. Miss
S. G. Stokes (The Genus Eriogonum, 36. 1936) recognizes
subsp. typicum and subsp. lappulaceum, considering other described
segregates as mere ecological variations. The varieties treated
in this paper, except var. lappulaceum would be included by her
in subsp. typicum. Since these plants as observed in the field
do not behave as ecological variants, I cannot agree with her
treatment.

KEY TO THE VARIETIES OF ErR1I0GONUM ABERTIANUM

Outer sepals obovate, almost truncate at top....... 7. EH. Abertianum
var. lappulaceum
Outer sepals nearly circular, broadest in the middle.

Plant paniculate above; bracts inconspicuous, up-
per part of plant not leafy.
Peduncles mostly under 0.5 mm. long; outer
sepals dark reddish, 2 mm. broad or nar-
TOWER. 52a 0y eels eh a a ete 2. H. Abertianum
var. ruberrimum
Peduncles mostly over 0.5 mm. long, outer sepals
only slightly reddish, 2.5 mm. broad or wider. 1. EH. Abertianum
var. neomexicanum
Plant not paniculate above, peduncles of involucres
axillary, plant leafy or conspicuously bracteate
to top.
Peduncles mostly under 1 cm. long, heads usually
under 1 cm. across, plant bracteate above.... 5. EH. Abertianum
var. bracteatum
Peduncles mostly over 1 cm. long, heads usually
over 1 cm. across, plant bracteate or folia-
ceous above.
Leaves reduced to lanceolate bracts above,
basal leaves attenuate at base, their petioles
2:¢em. long or under. +0 fe eee eee 6. EH. Abertianum
var. Gillespiet
Leaves not much reduced above, basal leaves
usually truncate or cordate at base,
petioles of basal leaves usually over 2 cm.
long.
Plants branching at base, usually floriferous
to base; outer sepals 3.5-4.5 mm. broad,
slightly longer than broad ............. 4, EH. Abertianum
var. cyclosepalum
Plants branching dichotomously or trichoto-
mously somewhat above base, floriferous
above only; outer sepals 2.5-3.5 mm. broad,
slightly broader than long ........:.... 3. EH. Abertianum
var. villosum

1938 ] FOSBERG: ERIOGONUM ABERTIANUM 191

1. Er1oconum ABERTIANUM Torr. var. NEOMEXICANUM Gandoger,

in Compt. Rend. Soc. Bot. Belg. 42: 196. 1906. LE. pinetorum

Greene, Muhlenbergia 6: 38. 1910.
Erect plants, soft villous but not conspicuously whitish except

on young parts, seldom branched at base but usually repeatedly

dichotomously or trichotomously branched above into a definite
panicle, peduncles and upper branches usually filiform; petioles

. 1-2 cm. long, leaves near base broadly oblong-ovate to narrowly

ovate, obtuse to acute at apex, truncate to attenuate at base, re-
duced above to small linear bracts 5 mm. or less long in the
paniculate inflorescence; peduncles 0.5—3 cm. long, filiform, usu-
ally 1 at a node, surmounted by a single turbinate involucre,
2-2.5 mm. high, 8 mm. in diameter with lanceolate lobes usually
2.5-4 mm. long; flowers rather numerous, exserted on fine fili-
form pedicels, the rather spherical inflorescence 5-10 mm. in
diameter; outer sepals 2.5-3 mm. broad, circular or slightly
longer than broad, membranous-scarious, yellowish-white, tinged

~ with red.

Specimens examined. Kansas. Arkansas River, Pawnee Co.,
July 17, 1846, Lt. Abert (type, New York Bot. Gard.). Arizona.
Tucson, Thornber 196; Tucson, May 10, 1896, J. W. Toumey; Out-
law Cafion, Chiricahua Mts., Goodding 2373; Pearce, Cochise Co.,
Aug. 19, 1910, W. W. Jones; Antelope, C. A. Purpus 52; Clifton,
A. Davidson 445. New Mexico. Rio Grande, Sierra Co., Oct. 14,
1846, Lt. Emory (cotype). Dona Ana County: vicinity of Pyra-
mid (Bishop’s Cap) Peak, Mesilla Valley, south end of the Organ
Mts., F. R. Fosberg 83300, S3487, S3550, S3653, S3700, S3744,

| §3790, 83794, 83801, 84047; mesa west of the Organ Mts., Oct.

17, 1908, E. O. Wooton; Organ Mts., E. O. Wooton 427 (type of
var. neomezicanum) ; Mogollon Mts., Sierra Co., H. L. Rusby 359.

2. Eriogonum ABERTIANUM Torr. var. RUBERRIMUM Gandoger,
in Compt. Rend. Soc. Bot. Belg. 42: 196. 1906.

Similar to var. neomexicanum but with peduncles mostly under
0.5 cm. long; flowers just as numerous in the involucre, but not
so far exserted, heads very compact, 5-6 mm. in diameter; outer
sepals 1.5—2 mm. broad, nearly circular, dark reddish in color.

Specimens examined. Mexico. Near Casas Grandes, Chi-
huahua, C. H. T. Townsend & C. M. Barker 369 (type).

8. Erroconum ABERTIANUM Torr. var. villosum var. nov.

Planta erecta alba-villosa ad basin versus simplex; pedunculi
robusti in axillis superioribus solitarii, 2-6 cm. longi; sepala
exteriora quam longa latiora.

Erect plant, up to 4 dm. tall, more or less white villous, more
so in young plants, stem usually not branched at base, dichoto-
mously or trichotomously (often several times) branched above,
floriferous ordinarily at upper nodes only, leafy to the top, the

192 MADRONO [Vol. 4

leaves somewhat smaller above, but not conspicuously reduced;
leaves oblong-ovate, acute to obtuse at apex, truncate or at least
abruptly contracted at the base; petioles 2-3 cm. long on lower
leaves, becoming shorter above to about 0.5 cm; peduncles solitary
in the upper axils, rather robust, usually 2-6 cm. long (very short
in material collected at Douglas, Ariz., by L. N. Goodding, May
22, 1907); involucre solitary, 3 mm. high, 3 mm. in diameter,
turbinate, with ovate-lanceolate lobes about as long as the tube;
flowers many, exserted on rather long pedicels, the spherical in-
florescence 1—1.5 cm. in diameter; outer sepals 2.5-3.5 mm.
broad, nearly circular, slightly broader than long, with a narrow
sinus at the base, membranous-scarious.

Specimens examined. Arizona. Road to Soldier's Camp,
Santa Catalina Mts., alt. 6000 ft., G. J. Goodman & C. L. Hitchcock
1263 (type, Herb. Univ. Calif. no. 426,460); Douglas, ZL. N.
Goodding 2265 (low plant, very villous, peduncles mostly short,
possibly quite young) ; Lowell, W. F. Parish 233 (in habit resem-
bling var. cyclosepalum) ; Chiricahua Mts., Sept., 1881, J. G. Lem-
mon and wife; Tucson, May 10, 1896, J. W. Toumey; Nogales, May
24, 1892, T. S. Brandegee; Douglas, May 17, 1915, Carlson; Ox
Bow Hill, Apache Trail, Eastwood 17524. New Mexico. Mim-
bres River, Grant Co., July 1, 1904, alt. 5500 ft., O. B. Metcalfe
1057; Mogollon Creek, Mogollon Mts., Socorro Co., alt. 8000 ft.,
O. B. Metcalfe 234; Deming, Grant Co., alt. 4400 ft., J. W. Gil-
lespie 53821. Texas. El Paso, April 18, 1884, M. E. Jones.
Mexico. San Requis, Lower Calif., May 2, 1889, T. S. Brandegee;
Dry Mts., east of Rio San Miguel, Chihuahua, C. V. Hartman 654

(last two collections extremely villous).

4. Ertoconum ABertianum Torr. var. cyclosepalum (Greene)
comb. nov. LE. cyclosepalum Greene, Muhlenbergia 6: 1. 1910.

Very similar to var. villosum in general appearance and in
pubescence but branched at base, depressed in habit, 1 dm. or.
less tall, floriferous to the base; peduncles 1-3 cm. long; invo-
lucres turbinate, the tube 2 mm. high, 3 mm. in diameter, the
lobes very large, more or less spatulate with rounded apex, 6—10
mm. long, 1-2 mm. broad; flowers many, the cluster 1—-1.5 cm.
in diameter; outer sepals 3.5—4 mm. broad, nearly circular but
slightly longer than broad, membranous-scarious, yellow tinged
with red.

Specimens examined. New Mexico. Mesa west of the Organ
Mts., Dona Ana Co., May, 1905, FE. O. Wooton; Silver City, East-
wood 8419. Texas. El Paso, M. E. Jones 3738; Fort Bliss, April
22,1915, Carlson. Mexico. Chihuahua, April, Dr. Parry; Janos
[Janas?] March, Dr. Parry; San Luis Potosi, J. G. Schaffner 2178.

5. Erroconum AsBertianum Torr. var. bracteatum var. nov.
Planta basi ramulosa, viridis; ramuli adscendentes pluriramu-

1938] FOSBERG: ERIOGONUM ABERTIANUM 193

losi saepe ad basin floriferi; folia parva supra reducta; pedunculi
axillares, 5 mm. longi; involucra parva 2 mm. lata altaque, lobis
4-6 mm. longis; sepala exteriora quam longa latiora.

Plant branched at the base, soft villous, green in color,
branches ascending, many times dichotomously or trichotomously
branched, floriferous often almost to base, forming usually a
small rounded plant, ordinarily not more than 2 dm. tall, but
when in very favorable localities reaching a height of 4 dm.;
leaves small, the lower short-petioled, ovate, usually obtuse at
apex, attenuate at base, rapidly reduced to narrowly lanceolate,
sessile bracts above, these seldom over 1 ecm. long, but quite
abundant; peduncles mostly about 5 mm. long, in axils from top
often practically to base of plant; involucre small, turbinate, 2
mm. high, 2 mm. in diameter, with oblong-lanceolate lobes 4—6
mm. long, few-flowered; flowers exserted on rather long pedicels,
the spherical inflorescence 5-10 mm. in diameter; outer sepals
3—3.5 mm. broad, nearly circular, slightly broader than long, with
a narrow sinus at the base, membranous-scarious, yellow, tinged
with red.

Specimens examined. New Mexico. Near San Antonio, So-
corro Co., Ferris & Duncan 2309; vicinity of Pyramid (Bishop’s
Cap) Peak, near south end of Organ Mts., Dona Ana Co., F. R.
Fosberg S3244, S8307, S3486 (type, Herb. Los Angeles Mus.),
S3641, S3654, S3798. Texas. Gravelly mesa north of Chisos
Mts., Brewster Co., J. A. Moore & J. A. Steyermark 3267; Fort
Davis, Davis Mts., Jeff Davis Co., Ferris § Duncan, 2641. Local-
ity uncertain. “Near San Diego (Vall. of R. Grande), August,
Bigelow.

6. Errogonum ABERTIANUM Torr. var. Gillespiei var. nov.

Planta var. bracteato similis sed diffusior; pedunculi 1—2 cm.
longi; involucra multiflora 3 mm. alta, 3-4 mm. lata.

Similar to var. bracteatum, but plant more diffuse; peduncles
mostly longer, 1-2 cm. long; involucres larger, many-flowered,
3 mm. high, 3-4 mm. in diameter, lobes 2.5—4.5 mm. long; inflo-
rescence 1—1.5 cm. in diameter, outer sepals apparently not yel-
lowish, as in var. bracteatum, but whitish, tinged with red, midrib
very noticeable, dark red.

Specimens examined. Arizona. Apache Gap, Pinal Co., 2500
ft., J. W. Gillespie 8797 (type, Herb. Univ. Calif. no. 489,490).

7. Eriogonum Asertianum Torr. var. lappulaceum (Greene)
comb. nov. EE. lappulaceum Greene, Muhlenbergia 6: 2. 1910.
E. Abertianum subsp. lappulaceum Stokes, Gen. Eriog. 37. 1936.

Erect or ascending, 1.5 dm. high, softly villous but not hoary,
branches few, alternate, in no degree cymose or paniculate;
leaves on lower undivided portion of stem rhombic-ovate to ellip-
tic, short petioled; involucres few-flowered, the lobes oblong,

194 MADRONO [Vol. 4

longer than the body, often equalling the flowers, outer sepals
obovate, almost truncately obtuse at apex (ew. char.).

I have not seen specimens of this variety but, judging from
the description, it belongs here. Greene cites only one collec-
tion: Camp Charlotte, Texas, 1889, Nealley.

University of Hawaii, Honolulu,
June, 1936.

AN UNDESCRIBED SPECIES OF VIOLA FROM UTAH

Mito S. BaKer

Viola Clauseniana sp. nov. Herba acaulescens; rhizoma sim-
plex, carnosum radices numerosas adventitias ferens; folia 3—5
em. lata, 4-5 cm. longa subtus venis sparsissime pilosa aliter |
glabra, prostrata deltoidea decurrentia crasse et acute serrata
apicibus obtusis; petioli 5-11 cm. longi alati; stipulae 1.5—2 em.
longae anguste lanceolatae, inconspicue glanduloso-dentatae ;
pedunculi 8-14 cm. longi crassi erecti; bracteolae 5-6 mm. longae
subulatae floribus propinquae; sepala 2-3 mm. lata 7-9 mm. longa
ovato-lancolata acutiuscula crasse nervata auriculis sparse pubes-
centibus; corolla lilacina petalis omnibus in parte inferiore pal-
lidioribus; petala superiora 7 mm. lata 15 mm. longa anguste
obovata, petala lateralia 7 mm. lata 15 mm. longa oblongo-ob-
ovata imberbia, petalum infimum 8 mm. latum 20 mm. longum
oblongo-spatulatum calcare obtuso complanatiusculo incluso;
staminum appendices anteriores ochroleucae posteriores 1.2 mm.
lata 3 mm. longae virides; pistillum 5 mm. longum; stylus capi-
tellatus ovario non flexus, tuba stigmatosa brevi nuda, foramine
diametro tertiam partem capituli; flores cleistogami complures;
capsula 6—7 mm. lata 8-10 mm. longa oblonga apice truncato;
semina 1.2 mm. lata 2 mm. longa nigra minute scabrida, pondere
1 mg.; caruncula subterminalis seminis dimidio brevior.

Acaulescent; rootstock simple, fleshy, giving rise to numerous
adventitious roots; leaves prostrate forming a rosette which may
reach 28 cm. in diameter; glabrous except for occasional hairs
along the veins on the ventral surface of the leaves; leaf blades
conspicuously deltoid, decurrent on the petiole, coarsely and
sharply serrate, obtuse at apex, 3 to 5 cm. wide, 4 to 5 cm. long,
on winged petioles 5 to 11 cm. in length; stipules narrowly lance-
olate, faintly glandular-toothed, 1.5 to 2 em. long; peduncles un-
usually stout, erect 8 to 14 cm. high, bractlets subulate, near the
flower 5 to 6 mm. long; sepals conspicuously nerved, slightly
pubescent on the auricles, ovate-lanceolate, somewhat acute, 2 to
3 mm. wide, 7 to 9 mm. long; corolla light violet with a lighter
center, 2.5 cm. in diameter; upper petals narrowly obovate 7 mm.
wide, 15 mm. long; lateral petals oblong-obovate, wholly beard-
less, 7 mm. wide, 15 mm. long; spur petal oblong-spatulate 8 mm.

1938] BAKER: VIOLA 195

wide at end and 20 mm. to end of broad somewhat flattened ob-
tuse spur; anterior appendages of stamens tan colored, closely
enclosing style, posterior appendages green, 1.2 mm. wide, 3 mm.
long; pistil 5 mm. long, style without flexure at ovary, capitate,
naked, stigmatic tube short, foramen one-third diameter of head;
cleistogamous flowers abundant after early spring, producing
most of the seeds; capsule oblong, truncate at apex, 6 to 7 mm.
wide, 8 to 10 mm. long; seeds minutely roughened, nearly black,
1.2 mm. wide, 2 mm. long, weight 1 mg., caruncle latero-terminal,
extending about one-half the length of the seed.

Type: Zion National Park, Utah, July 5, 1936, M. S. Baker
8438 (Herb. Univ. Calif. no. 575768). This species was collected
at the base of Weeping Rock on the south side of the canyon
where sunlight seldom, if ever, reaches. In competition with
rather rank vegetation, the leaves of some of the plants are erect
and very tall but in the open they spread out without reaching a
great height. Transplanted to the garden at Kenwood, Cali-
fornia, the plants grow vigorously producing an enormous num-
ber of seeds, mainly from cleistogamous flowers which develop
capsules until growth is stopped by winter cold. The area in
which this species grows is very small, scarcely one hundred feet
in diameter (as remembered), densely covered with shrubs, trees
and herbaceous plants. The only known plants grow near a trail
used by thousands of tourists and, unless measures are taken to
protect them, are in great danger of extermination. Mr. K. E.
Weight, naturalist during the summer of 1936, reports that this
species does not occur elsewhere in the Park.

Viola Clauseniana belongs in section Nomimium Ging, subsec-
tion Plagiostigma Godr. group Boreali-Americanae Becker, series
Cucullatae Gersh. It is sharply distinguished from any other
known member of this series by the beardless petals, the deltoid
leaf outline and the prostrate habit of the leaves. Viola nephro-
phylla Greene is its nearest known relative.

At a higher elevation (9000 feet) at Navaho Lake, only a few
miles distant, was found another member of this group, Viola
arizonica Greene. ‘This species however is much more closely
related to V. nephrophylla than to V. Clauseniana. At a lower ele-
vation and only about thirty miles distant by airline, the writer
collected V. nephrophylla in typical form. This locality is seven
miles north of Kanab, Utah, along the highway to the north rim
of Grand Canyon. These plants showed not the slightest vari-
ation toward the species under discussion. Under such circum-
stances one is forced to the conclusion that V. Clauseniana is a
relict of an earlier age, not closely related to any living species,
which has survived at this one spot in Zion National Park.

This violet is named in honor of: Dr. Jens Clausen, cyto-
geneticist of the Carnegie Institution of Washington, Stanford

196 MADRONO [Vol. 4

University, California, who has been most helpful in solving
many problems connected with our western violets.

Santa Rosa Junior College,
Santa Rosa, California,
December 6, 1937.

REVIEWS

Botanical Studies in the Uinta Basin of Utah and Colorado. By
Epwarp H. Grauam. Annals of the Carnegie Museum. Volume
XXVI. Pp. 1-432. Pl. I-XIII. Carnegie Museum, Pittsburgh,
Pennsylvania. 19387.

This comprehensive study of one of the little known natural
areas of the western United States represents the results of three
Carnegie Museum expeditions to the Uinta Basin in the summers
of 1981, 1933, and 1935. In this work 1104 species of vascular
plants are recorded from the Basin, an area of 12,000 square
miles. Five new species and two new varieties are included. The
annotated list of species is based upon collections by the author
of some 8500 numbers and a few additional collections by others.
Most of these specimens are deposited in the herbarium of the
Carnegie Museum. In addition to the annotated list there are
discussions of the history of exploration in the Basin, the physiog-
raphy and climate of the area, the altitudinal vegetation zones
and plant communities. The geographic affinity of the flora and
endemism are discussed briefly. A list is given of range exten-
sions. The bibliography of the region lists 93 publications.

This study shows a minute attention to detail in its prepara-
tion; most of the identifications were checked by specialists; and
the entire volume shows the author’s intimate knowledge of the
botany of the region. Such a complete survey of a natural
geographic area is a definite contribution to our knowledge of
western flora—Mu1.prep E. Maruias.

Illustrated Dictionary of Botanical Terms. Excerpt from “An
Introduction to Botany” by Joun Linptey, 1848, containing pages
319, 346-383. Reprinted by Atice Eastwoop, California Acad-
emy of Sciences, San Francisco, with the cooperation of the San
Francisco Garden Club. 1988. Paper. $.50.

Originally published in 1848 this illustrated glossary of
botanical terms with Latin equivalents used in the description and
naming of plants has proved most useful to professional and
amateur botanists. The reprinting of the pages of this glossary
in a convenient pamphlet is a valuable contribution. Copies are
obtainable from Miss Eastwood.—Miuxprep E. Marutias.

The California Salvias. A Review of Salvia, Section Audibertia.
By Cart Epuine. Annals of the Missouri Botanical Garden.
Volume XXV, Number 1. Pp. 95-188 with 19 plates and 14 text —
figures. St. Louis, February, 1938.

1938 REVIEWS 197

In Audibertia, a section of the genus Salvia confined to arid
southwestern North America, Dr. Epling recognizes eighteen
species which he treats under five sections. Three of these,
Greeneostachys, Jepsonia, and Parishiella, are new; two, Echino-
sphace and Pycnosphace, reduced from the sectional rank given
by Bentham. One new subspecies, Salvia carnosa subsp. Gilmani,
and twelve new hybrids are described. The distribution of each
species is shown by a map and the habit and floral characters of
each beautifully illustrated by a full page plate. The introduc-
tion includes a discussion of the distribution and habitats of the
species of the section and their relation to the shrub formations
of the Colorado Desert and the coastal plain and foothill region
of southern and central California. Because of the interchang-
ing of blocks of characters in the species throughout the section
a hybrid origin is postulated for the group; the species show no
evidence of a monophyletic origin. The subsections are segre-
gated mainly upon stamineal and other floral characters; habit
and leaf characters have also proved useful. This conservative
and competent revision is the result of ten years of field work
and herbarium study on the part of the author, and is a worthy
successor to his many valuable contributions to the taxonomy of
the Labiatae. It is one of seven important botanical papers ap-
pearing in the current issue of the “Annals” which is dedicated
to Dr. Jesse More Greenman, Curator of the Herbarium of the
Missouri Botanical Garden, and which was prepared in honor
of his seventieth birthday by a group of his former students.—
EK. Crum.

A Revision of the Genus Lomatium. By Mitprep E. Marutas.
Annals of the Missouri Botanical Garden. Volume XXV, Num-
ber 1. Pp. 225-297. St. Louis, February, 1938.

This largest and perhaps most difficult of West American
umbelliferous genera is interpreted as consisting of 63 species
and 20 varieties. Two species and two varieties are described
as new. <A conservative point of view has been maintained
throughout the treatment, despite the superabundance of names
and the frequent paucity of available specimens of some units.
The appearance of the publication, in itself, should stimulate col-
lectors working in regions immediately concerned to obtain ade-
quate material for the solution of the few problems and the filling
of occasional gaps in distribution which still remain.

One notes, gratefully, that the key does not rely unduly upon
the number of oil tubes in the fruit for the separation of species
and varieties, but that vegetative, floral and habital characters as
well as geographical ranges are used to facilitate identification.
The species are clear-cut and in the great majority of cases pre-
sent a logical distribution pattern which conforms with that in
other large genera whose taxonomy and distribution have been

198 MADRONO [Vol. 4

studied. Species of narrowly restricted range occur only in such
areas as are suspected, on other evidence, of possessing endemic
floras. Many of the “‘one-specimen species” have been found to
fit snugly into groups of wider occurrence.

In previous papers, the author has shown a preference for the
generic name Cogswellia Spreng., both because of the question-
able identity of the type species of Lomatium Raf. (which ante-
dates it by one year) and the close similarity of the latter name
to Lomatia R. Br. of the Proteaceae. The present manuscript
was written with the intention of retaining Cogswellia, but a
hurried poll of available authorities on nomenclature, taken at
the suggestion of the editor, necessitated a last-minute substitu-
tion of Lomatium. Now that all the transfers have been made
to Lomatium, it is to be hoped that this interpretation of the
International Rules of Nomenclature will be upheld.

One can readily recognize in the treatment the background
of the author’s extensive knowledge of the Umbelliferae and her
unusually broad field experience with the family. Because of the
thoroughness and practicality of the treatment, one awaits with
interest the appearance of revisions of other troublesome genera
of this family.—L. Constance.

Plants of Zion National Park. By Cuirrorp Presnatt and
PautinE Meap Patraw. Zion-Bryce Museum Bulletin No. 1.
Zion-Bryce Natural History Association in cooperation with the
National Park Service. June, 1937. Pp. 1-69 with 15 plates and
lbitext fourées;: Paper: $.50.

A brief synoptical treatment of the common flowering plants
and ferns of Zion National Park. The common names are em-
phasized in accordance with the intended popular appeal. The
work consists of a list of over five hundred of the known species
of the Park. In most cases a brief statement of characters or
habitat accompanies the names. There are no formal descriptions
or keys. The illustrations are line drawings and photographs.
The printing is by the offset process——HersBert L. Mason.

Die Bedeutung der Polyploidie fiir die Verbreitung der Angio-
spermen, erldutert an den Arten Schleswig-Holsteins, mit Ausblicken
auf andere Florengebiets. By G. Tiscuier. Bot. Jahrb. Band
LXVII, Heft 1. Pp. 1-36. Leipzig. 1935.

The chromosome numbers of 66.7 per cent of the species of
angiosperms of Schleswig-Holstein are recorded, and 44.1 per |
cent are found to be polyploid. Families rich in polyploids are
the Polygonaceae, Rosaceae, Malvaceae, Rubiaceae, Gramineae,
and Cyperaceae, while the Leguminosae and Umbelliferae have
relatively few such species. Of the circumpolar types found in
this province, 60 per cent are polyploid, while of those of a more
southerly range than the area investigated, only 27.1 per cent

1938 | PROCEEDINGS 199

are of this type. In Iceland and the Faroe Islands the percent-
age of polyploids is higher than in Schleswig-Holstein; in Sicily
it is lower. The majority of the polyploid species of Iceland
are wide ranging types rather than species endemic to the island
or to the Arctic flora. In the flora of a restricted locality, such
as an estuary and its borders, many polyploids are found among
the species occupying that as well as other habitats, and fewer
among the species peculiar to that habitat. The greater varia-
bility of polyploids is responsible for this condition, since it
broadens their range of tolerance to different habitats.—G. L.
STEBBINS, JR.

NOTES AND NEWS

CEANOTHUS THYRSIFLORUS: EXTENSION OF RANGE. A considerable
stand of this species, previously not known to occur south of
Monterey County, was observed by Mr. Maunsell Van Rensselaer
on a north slope of upper Canada Honda, southwest of Lompoc,
Santa Barbara County, California (February 27, 1938, M. Van

_ Rensselaer, Herb. Univ. Calif.). The shrubs were from twenty

to twenty-five feet in height; some with a branch spread of over
thirty feet—H. E. McMinn.

The Wild Flower Show of Santa Rosa Junior College which is
held biennially will open on Sunday, May 15, 1938 in the Science
Building of the College at 1:00 P.M. In 1936 Professor Milo S.
Baker and his assistants had representatives of over one thousand
native Californian species on display.

The “Helen Stafford Thorne Ceanothus Fund” was recently
established by the western member clubs of the Garden Club of
America in honor of Mrs. Oakleigh Thorne who has been its
active representative on the Pacific Coast for many years. The
fund has been given to the Blaksley Botanic Garden to be used
in furthering the study of the genus Ceanothus. (Santa Barbara
Museum of Natural History Leaflet 138: 22. 1938.)

Dr. W. A. Setchell, Professor of Botany, Emeritus, Univer-
sity of California, Berkeley, has received notice of his election
as a Foreign Member of the Royal Society of Science and Arts,
of Gothenburg, Sweden. This honor comes as a result of studies
made in cooperation with Swedish botanists.

PROCEEDINGS OF THE CALIFORNIA
BOTANICAL SOCIETY

November 18, 19387. A meeting was held at 8:00 p. m. in
Room 2098, Life Sciences Building, University of California,
Berkeley. The president, Dr. F. W. Foxworthy, occupied the
chair. Lecture: “An American botanist in Europe” by Profes-

200 MADRONO [Vol. 4

sor H. E. McMinn, of the Department of Botany, Mills College,
California. Mr. McMinn recounted experiences during a recent
visit to botanical institutions of Sweden, Norway and other
European countries.

December 16, 1937. A meeting was held in Room 2093, Life
Sciences Building, University of California, Berkeley. The
president, Dr. F. W. Foxworthy, called for the report of the
nominating committee. Professor E. B. Babcock, chairman, sub-
mitted the following names: President, Professor H. E. McMinn,
Mills College, California; First Vice-President, Mrs. Viola Brain-
erd Baird, Berkeley, California; Second Vice-President, Mr.
Maunsell Van Rensselaer, Blaksley Botanic Garden, Santa Bar-
bara, California; Treasurer, Dr. David D. Keck, Carnegie Insti-
tution of Washington, Stanford University; Secretary, Miss Ethel
Crum, University of California, Berkeley. The business meeting
was followed by a lecture: ““A quest for western wild violets” by
Viola Brainerd Baird. Mrs. Baird, daughter of Ezra Brainerd,
the well known authority on North American violets, described
the haunts of the western species of the genus. Her lecture was
illustrated with a complete series of colored slides reproduced
from original paintings by F. Schuyler Mathews.

January 27, 1938. A meeting was held at 8:00 p. m. in
Room 2098, Life Sciences Building, University of California,
Berkeley. The president, Dr. F. W. Foxworthy, occupied the
chair. The officers nominated at the preceding meeting were
unanimously elected. Following the business meeting Dr. Lin-
coln Constance, of the University of California, Berkeley, lec-
tured on ““Endemism in the Pacific Northwest.”

The following fall and winter meetings have been held by the
Santa Barbara Branch of the California Botanical Society.

October 19, 1987. Lecture: “A quest for western wild
violets’ by Mrs. Viola Brainerd Baird, Berkeley.

November 16, 1987. Lecture: “‘The forest and. the trees”
by Dr. George J. Peirce, Department of Botany, Stanford Uni-
versity.

February 1, 1988. Lecture: ‘“‘Plant exploring in Mexico” by
Mr. Otis McAllister, for many years a resident of Mexico City.

March 12, 1988. A joint conference of officers of the two
branches of the California Botanical Society was held. The
parent organization was represented by Professor H. E. McMinn,
president, Dr. George J. Peirce, former president, and Dr. Her-
bert L. Mason, chairman of the editorial board of MaproNno.
President Maunsell Van Rensselaer and the board of control
represented the Santa Barbara Branch.

March 14, 1988. Lecture: ‘““Why an American botanist visits
European herbaria to study native American plants.” Professor

H. E. McMinn, Mills College, California.—E. Crum, Secretary.

Complete Your Files!

MADRONO

A West American Journal of Botany

“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges.

_ Volume I, 1916-1929. . . $5.00
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Single numbers....... O75

There is only a limited supply of Vol-
umes I and II.

Address all communications
and orders to:

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Stanford University, California

Silver fubilee Colebration
1913—1938

Twenty-fifth anniversary of the founding

of the

CALIFORNIA BOTANICAL SOCIETY
To Honor the Charter Members of the Society

April 23, 1938

Dinner at 6:15 P. M.
at the

Berkeley Women’s City Club

2315 Durant Ave., Berkeley, California

Price One Dollar
Per Plate

Send reservations with check to Dr. Lincoln Constance, Department of Botany
Life Sciences Building, University of California, Berkeley.

OLUME IV so AMEN INSTT oes NUMBER 7

Rig woh snl 3, {#7

C ee Tac eK RDI

MADRONO |

A WEST AMERICAN JOURNAL OF
BOTANY

4

: = :

Na a a

Contents
A CRANBERRY FROM THE TAHOE NatTIonaL Forest, William A. Dayton .... 201
TUDIES IN THE GENUS STREPTANTHUS Nutt. I. Two New SPECIES IN THE
Section Evcuista Nort., John L. Morrison .......... 0.00. cc ees 204

Tue Native CatirorNnian SPECIES OF THE GENUs Coreopsis L., Helen K.

A

Ce IEE TUM rt MN SN ie gaia | a yt io ele ici lg whale d wlanhlntelele ace eheield 209

Coton Vartation 1x DELPHINIUM CARDINALE Hoox., Charles O. Blodgett
| PAG OE NOME UTSE: cee ee ae cl A a occa a Ge 231
| ——- Guavucocarrum, a New GENUS IN THE CrucireRAE, Reed C. Rollins ....... 232

Reviews: Voegelin Tibatulabal Ethnography (Herbert L. Mason) ; Smith,
Species Lupinorum (Herbert L. Mason); Applegate, Plants of the

Lava Beds National Monument, California (Lincoln Constance) ..... 236
Notes anp News: Battarea phalloides (Dicks.) Pers. in Santa Barbara

etary rebar ewe ys MMe oN NS Ue a ue Cll wis | 238

PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY ................-205: 239

Published at Lime and Green Streets, Lancaster,
Pennsylvania

July, 1938

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Board of Editors

Dr. H. L. Mason, University of California, Berkeley, Chairman.

Dr. L. R. Asprams, Stanford University, California.

Dr. Lincotn Constance, University of California, Berkeley.

Dr. H. F. Copetanp, Sacramento Junior College, Sacramento, California.
Dr. A. W. Haupt, University of California at Los Angeles.

Business Manager—Dr. Davin D. Keck
Lime and Green Sts., Lancaster, Pennsylvania
or
Carnegie Institution of Washington
Stanford University, California

Entered as second-class matter October 1, 1935, at the post office at
Lancaster, Pa., under the act of March 3, 1879.

Established 1916. Published quarterly. Subscription Price $2.50 per year.
Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17,
complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num-
bers $0.75.

Papers up to 15 or 20 pages are acceptable. Longer contributions may
be accepted if the excess costs of printing and illustration are borne by the
contributor. Range extensions and similar notes will be published in con-
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Published at Lime and Green Streets, Lancaster,
Pennsylvania, for the

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Professor H. E. McMinn, Mills College, California. First Vice-
President: Mr. Maunsell Van Rensselaer, Blaksley Botanic Garden, Santa
Barbara, California. Second Vice-President: Mrs. Viola Brainerd Baird,
Berkeley, California. Treasurer, Dr. David D. Keck, Carnegie Institution of
Washington, Stanford University, California. Secretary: Miss Ethel Crum,
4004 Life Sciences Building, University of California, Berkeley.

Annual membership dues of the California Botanincal Society are $2.50,
$2.00 of which is for a year’s subscription to Madrofio. Dues should be
remitted to the Treasurer. General correspondence and applications for
membership should be addressed to the Secretary.

1938] DAYTON: CRANBERRY 201

A CRANBERRY FROM THE TAHOE NATIONAL FOREST
Wittiam A. Dayton

No botanical manual or other floral publication, so far as the
writer has ascertained, attributes cranberries to the California
flora. Mr. Leland S. Smith of the Supervisor’s staff of the Tahoe
National Forest, an indefatigable collector and observer of Sier-
ran plants, has collected two very interesting specimens of cran-
berry in Nevada County.

Oxycoccus macrocarpos' (Ait.) Pursh. Hydraulic digging, one
mile south of North Columbia, 3000 feet elevation, with pon-
derosa pine (western yellow pine), willows and sphagnum and
other mosses, November 14, 1936, L. S. Smith 2800 (United States
Forest Service serial no.73019), sterile specimen; bog, southwest
side of Columbia Hill diggings, 2900 feet elevation, July 17, 1937,
L.S. Smith 2800A (United States Forest Service serial no. 75678),
specimen in immature fruit, with some late flowers. Duplicates
of the above collections are deposited in Forest Service herbaria
at Nevada City and San Francisco, California, and at Washington,
DC:

In a memorandum of July 26, 19387, Mr. Smith writes:

This species evidently blooms earlier than I thought, as I found a large
amount of half-formed berries. Flowers no doubt appear around July 1 or
possibly some years, in June.

I have visited other similar sites and have not found any indication of this
species being present. So far as I can learn, this is the only place on the Forest
where it is found, and no one now living in the locality (North Columbia) knows
anything about it. It was only recently discovered, and it seems possible that
some cranberries were dropped or thrown into the water in the early days by
some miner, and became lodged in this pocket, at the end of the hydraulic min-
ing in this vicinity, which was in the early eighties.

Fruit was collected here, and made very fine jelly and sauce. The bog
is limited in size, and area of cranberry plants will not grow much unless seed
is carried to other bogs by rodents or birds.

This material was provisionally identified by Mr. Smith as
Vaccinium oxycoccos’ L. var. intermedium A. Gray. It seems natural
that, if any cranberry were to be found in California, it would be
this variety. However, in view of the rather long, narrowly
oblong, clearly revolute leaves, distinctly rounded and blunt at
both ends, the small but foliaceous and relatively broad bractlets
subtending the pedicels, and the large fruits, typically longer
than broad, the plant, is, in my judgment, the commonly culti-
vated cranberry, Oxycoccus macrocarpos (Ait.) Pursh (Vaccinium
macrocarpon Ait.), sometimes called big cranberry or American
cranberry. The so-called small (European) cranberry, Oxycoc-

Manprono, Vol. 4, pp. 201-240, July 1, 1938.
1 In this case the Latin ending -ws is usual in literature, but in view of the
original spelling the Greek termination -os should be used.

m Pye
we ?

202 MADRONO [Vol. 4

cus palustris Pers. (O. oxycoccos (L.) MacM., Vaccinium oxycoccos
L.) is also native in parts of our country and is occasionally cul-
tivated. Its smaller, round fruits are considered by many to
have a flavor superior to those of Oxycoccus macrocarpos. In our
Northwest there occurs a native variety of the small cranberry,
to which I have applied’? the name “‘western cranberry,’ O. palus-
tris var. intermedius (A. Gray) Howell (O. oxycoccos intermedius
(A. Gray) Piper, O. intermedius (A. Gray) Rydb., Vaccinium oxy-
coccos var. intermedium A. Gray). This differs from typical
forms of the species, chiefly in its coarser stems, blunter, broader
and less revolute leaves, larger fruit, and in a pronounced ten-
dency for the shoots to proliferate after flowering.

Herbarium material of these three cranberries is often badly
mixed and it is quite possible to match Mr. Smith’s Tahoe plants
with specimens in folders labeled Oxycoccus macrocarpos, O. palus-
tris or O. palustris var. intermedius. Perhaps the best generally
available key to these three forms is that by Rydberg (FI. Rocky
Mts. 646. 1917). Dr. Rydberg regarded O. intermedius as a
species, although, as its name implies, it appears to be intermedi-
ate between O. palustris and O. macrocarpos. It may be a natural
hybrid between those species, a problem which should be investi-
gated by geneticists and cytologists.

The generic separation of Oxycoccus from Vaccinium, which
dates from Tournefort, is, of course, a matter of taxonomic con-
cept. The writer prefers to maintain this separation as was done
by Dr. Frederick V. Coville. One can hardly forbear an expres-
sion here of sorrow and regret at the passing of Dr. Coville, our
foremost American student of Vacciniaceae.

Mr. Smith properly mentions the possibility that this cran-
berry had been introduced accidentally by man in this Cali-
fornia bog. The possibility of introduction by birds, such as
grouse, or by other animals should also be considered. I see
no reason, however, why some cranberry might not locally be
native “in the higher northern parts of the Sierra Nevada” of
California, as Brewer and Watson suggested (Bot. Calif. 1:
450. 1876) about sixty years ago. At any rate, Mr. Smith's
keen eye and enthusiasm has added another genus and species to
the known flora of California.

Range Forage Investigations,
Division of Range Research,
United States Forest Service,

Washington, D. C.,
November 24, 1937.

2 Dayton, W. A. Important western browse plants. U. S. Dept. Agric.
Mise. Publ. 101. 1931.

DAYTON:

SS

X . RYE,
a= >

ED

ae:
SS

CRANBERRY

Pirate XXX. Oxycoccus macrocarpos (Ait.) Pers.
specimen; B, immature fruiting specimen; C, flower; D, stamens; E, calyx and
style; F, G, immature fruits; H, portion of stem with leaves.

Miss Leta Hughey of the United States Forest Service from specimen collected
in Tahoe National Forest, California, by L. S. Smith (no. 2800A).

\yj,

¥.

WY

VY, Sf

Wy

P ¢
iy l
YS AS j

Ny WZ

Centimeters

ee ———————

A, late flowering

Drawing by

203

204 MADRONO [Vol. 4

STUDIES IN THE GENUS STREPTANTHUS NUTT.
I. TWO NEW SPECIES IN THE SECTION
EUCLISIA NUTT.

Joun L. Morrison

During the course of monographic studies in Streptanthus it
has become necessary to describe two hitherto unrecognized spe-
cies. The decision to recognize the following as new species was
based on differences in the structure, shape, size and color of
organs as well as upon geographic distribution. The fused fila-
ments of the upper and lower pairs of stamens, the obtuse, crisped
petal blade, the non-bracteate inflorescence, and the annual dura-
tion of the plants indicate the close relationship betwen these
species and the others at present included in Section Euclisia.

Streptanthus batrachopus sp. nov. Herba annua omnino
glabra glaucaque maculata; caules erecti simplices vel superne
ramosi, 4.0-18.0 cm. alti; folia pauca, plerumque basalia crassa,
subtus purpurea, supra fulva vel purpureo-maculata, 0.3-3.2 cm.
longa, 0.2-1.0 cm. lata; folia inferiora, spatulato-obovata vel
oblonga, saliente lobata, petiolis longis; folia superiora sessilia
auriculata amplexicaulia, lineari-lanceolata vel oblonga, sub-
integra; flores erecti, 0.4-0.5 cm. longi; sepala 0.4 cm. longa,
viridia purpureave, ovata, basi carinata, apicibus patentibus acutis,
marginibus hyalinis, saepe rubescentibus; petala valde exserta
alba, venis mediis purpureis, lineari-lanceolata spatulatave, acuta

0.6-0.7 cm. longa; stamina triseriata, superiorum filamentis 0.6 —

cm. longis ad apicem connatis, antheris reductis, inferiorum fila-
mentis 0.5 cm. longis, usque ad 0.2—0.3 cm. connatis, antheris
longioribus, lateralium filamentis 0.3 cm. longis, liberis, approxi-
matis, antheris longissimis; siliquae 2.5-3.0 cm. longae, erectae,

leviter torulosae, falcatae, apicibus patentibus, virides purpureo- ~

maculatae; stylus 0.1 cm. longus; stigma integrum; pedicelli 0.2—
0.3 cm. longi, adscendentes ; semina fulva, striato-reticulata, alata,
0.2 cm. longa; cotyledones accumbentes.

Annual; stem erect, simple or branched above, entire plant
glabrous and glaucous, mottled, 4.0-18.0 cm. high; leaves few,
mostly basal, thick, purple beneath and brown or purple spotted
above, 0.8-3.2 cm. long; 0.2-1.0 cm. wide; lower leaves long-
petioled, spatulate-obovate to oblong, saliently-lobed; upper
leaves sessile, auriculate-clasping, linear-lanceolate to oblong,
sub-entire; flowers erect, 0.4-0.5 cm. long; sepals green or purple,
ovate and keeled at base, narrowed above, tips spreading, mar-
gins hyaline or reddish, 0.4 cm. long; petals well exserted, white
with purple midvein, linear-lanceolate to spatulate, acute, 0.6—
0.7 cm. long; stamens in three pairs, anthers saggitate; upper
pair 0.6 cm. long, filaments connate to the apex, anthers reduced;

1938 ] MORRISON: STREPTANTHUS 205

lower pair 0.5 cm. long, filaments connate for about half to two-
thirds their length, anthers longer; lateral pair free, approxi-
mate, 0.38 cm. long, anthers longest; silique 2.5-3.0 cm. long,
erect, falcate-spreading, slightly torulose, green, spotted with
purple, style about 0.1 cm. long, stigma entire; pedicels 0.2—0.3
em. long, ascending; seeds brown, striate-reticulate, winged,
about 0.2 cm. long, cotyledons accumbent.

Serpentine outcrops in Marin County, California, Mt. Tamal-
pais, North Side Trail, one-fourth mile northeast of Rifle Camp,
June 8, 1987, J. L. Morrison 2493 (type, Herb. Univ. Calif. no.
575762); May 238, 1937, J. L. and F. L. Morrison 2368 (Herb.
Univ. Calif.); Big Carson Ridge, 5.5 miles northwest of Mt.
Tamalpais, between Pine Mountain and San Geronimo Ridge,
May 30, 1987, J. L. Morrison 2440 (Herb. Univ. Calif.).

Streptanthus batrachopus is known at present only from two
outcrops of serpentine in Marin County. The plants are fairly
abundant locally, growing in serpentine talus. Associated plants
include: Allium falcifolium Hook. & Arn., Cheilanthes siliquosa
Maxon, Eriogonum vimineum Dougl. var. caninum Greene, Strept-
anthus glandulosus Hook. var. pulchellus (Greene) Jepson, Epi-
lobium minutum Lindl., Arenaria Douglasi Fenzl, Ceanothus Jepsoni
Greene, Arctostaphylos montana Eastw., Quercus durata Jepson and
Cupressus Sargent Jepson. Careful exploration of the many
other serpentine outcrops in the Mt. Tamalpais area has so far
failed to extend the range of this species. This name was chosen
because of the resemblance of the leaves of this species to a
frog’s foot. |

This species seems to be most closely related to Streptanthus
Breweri Gray, a serpentine endemic ranging in the inner and
middle Coast Ranges from Tehama County to San Benito County.
The key below indicates the differences between the two species:
Leaves broadly ovate, remotely denticulate to entire,

sessile or with a very short petiole, auriculate,

concolorous above at maturity, average lJength

4.1 cm.; flowers many, 0.6-0.8 cm. long; stigma

Sesciles SeeGsS WINGIESS fee le en es ale Streptanthus Breweri
Leaves spatulate-obovate, saliently few-lobed, long-

petioled, anthocyanous below, greenish-yellow

above, mottled with brown, purple or red; aver-

age length 1.2 cm.; flowers few, 0.4-0.6 cm. long;
style 0.1 cm. or more in length; seeds winged ... Streptanthus batrachopus

Streptanthus callistus sp. nov. Herba annua, compacta,
-omnino sparse hispida; caules erecti, 3.0—6.0 cm. alti, simplices
vel plerumque ramosissimi, ramis divaricatis, apicibus adscenden-
tibus; folia basalia subsessilia, oblongo-orbiculata, crasse dentata,
0.2—-0.6 cm. longa; folia superiora orbiculato-obovata vel oblonga,
basi lata, auriculata, amplexicaulia, dentata, 0.5-1.5 cm. longa,
0.4—-1.4 em. lata, saepe rubescentia, pilis rigidis, hyalinis, attenu-
atis, utrinque sparse hispida; racemi flores terminales steriles re-

206 MADRONO [Vol. 4

ducti, calycibus elongatis, hispidis, partibus ceteris obsolescenti-
bus; flores fertiles numerosi; pedicelli, breves, crassi, adscen-
dentes; sepala viridia, hispida, ovato-lanceolata, acuta, saccata,
dorsaliter carinata, 0.5 cm. longa; petala purpurea, conspicue,
venata, spatulato-oblanceolata, obtusa, 1.0 em. longa, marginibus
undulatis albidis, laminis 5 cm. latis; stamina triseriata, antheris
leviter sagittatis, superiorum filamentis 0.6 cm. longis fere usque
ad apicem connatis, antheris reductis, inferiorum filamentis 0.4—
0.5 cm. longis, usque ad 0.2—0.3 cm. connatis, antheris longiori-
bus, laterialum filamentis 0.38 cm. longis, liberis, approximatis,
antheris longissimis; siliquae 1.5—2.0 cm. longae, teretes, erectae,
incurvatae, attenuatae, valvis conspicue uninervatis, pilis com-_
planatis, hispidis; septorum cellulae tortuosae; semina olivacea,
globosa, non alata; cotyledones accumbentes.

Low compact annual, sparingly hispid throughout; stem erect,
simple or mostly much branched, branches at right angles to the
main stem, curving upwards, 3.0—6.0 cm. high; basal leaves sub-
sessile, 0.2-0.6 cm. long, oblong-orbicular, coarsely dentate;
upper leaves sessile, 0.5—1.5 cm. long, 0.4—-1.4 cm. wide, dentate,
often reddish, very sparsely hispid above and below with stiff,
hyaline, tapered trichomes, orbicular-obovate to oblong, base
broad, auriculate-clasping; raceme terminated by sterile flowers
reduced to elongated, hispid calyces, with petals, stamens and
pistil rudimentary; fertile flowers numerous on short, stout, his-
pid, ascending pedicels; sepals 0.5 cm. long; green, hispid, ovate-
lanceolate, acute, keeled, saccate; petals mulberry purple’ to
cotinga purple with prominent veins, 1.0 cm. long, spatulate-ob-
lanceolate, obtuse, margins undulate, whitish, lamina 0.5 cm.
wide; stamens in three pairs, anthers slightly saggitate; upper
filaments 0.6 cm. long, connate nearly to the apex, anthers re-
duced; lower filaments 0.4—-0.5 cm. long, connate for 0.2—0.3 cm.
anthers longer; lateral filaments 0.3 cm. long, free, approximate,
anthers longest, free tips of the connate filaments mulberry
purple; silique 1.5—2.0 cm. long, terete, erect, incurved, tapering,
valves with a strong midvein, hispid with flattened trichomes,
cells of the septum all tortuous; seeds greenish-brown, orbicular,
not flattened, wingless, cotyledons accumbent.

1 Color Standards and Nomenclature. Robert Ridgway. Washington, D. C.
Published by the author. 1912.

EXPLANATION OF THE Ficures. PLATe XXXII

Pirate XXXI._ Figs. 1-10, Streptanthus callistus Morrison: 1 habit; 2 ter-
minal cluster of sterile flowers; 3 flower; 4 petal; 5 seed; 6 flower; 7 androe-
cium and gynoecium; 8 leaf; 9 silique; JO sepal. Figs. 11-15, Streptanthus
hispidus Gray: 11 seed; 12 habit; 73 leaf; 74 androecium and gynoecium; 15
flower. Figs. 16-19, Streptanthus insignis Jepson: 16 seed; 17 leaf; 18 flower;
19 silique. Figs. 20-28, Streptanthus batrachopus Morrison: 20 silique; 21
habit; 22 sepal; 23 flower; 24 petal; 25 androecium and gynoecium; 26 leaf;
27 seed; 28 leaf. Figs. 29-31, Streptanthus Breweri Gray: 29 silique; 30 leaf;
31 seed.

1938 ] MORRISON: STREPTANTHUS

Pirate XXXI. CoMmMPaARATIVE DRAWINGS OF
(See explanation of figures on page 206.)

FIVE

SPECIES

OF

STREPTANTHUS,

208 MADRONO [Vol. 4

Arroyo Bayo, Mt. Hamilton Range, Santa Clara County, Cali-
fornia, 6.8 miles east southeast of Isabel Creek on the road to
San Antonio Valley, elevation 2000 feet, May 5, 1935, C. W. and
H. K. Sharsmith 3074 (type, Herb. Univ. Calif. no. 575766; ‘iso-
type, Gray Herbarium) ; May 138, 1987, D. D. Keck and J. Clausen
4541 (Herb. Carnegie Inst. Wash., Stanford Univ.); April 27,
1938, J. L. Morrison and A. Carter 3019 (Herb. Univ. Calif.).

Streptanthus callistus was discovered by Herbert L. Mason,
Carl W. and Helen K. Sharsmith in 1935. In 19387 a few plants
were collected by David D. Keck and Jens Clausen who report
it as “‘very rare.” The writer collected mature siliques and seed
in this area in October, 1937, and on April 27, 1938, the plants
were found to be abundant in the area. This very narrow en-
demic, known only from the type locality, occupies low, south-
facing knolls, where the soil is very loose and dry. Associated
with it are found: Malacothrix obtusa Benth., Eriogonum sp.,
Chaenactis glabriuscula DC., Mimulus Bolanderit Gray, Lessingia
germanorum Cham., Linanthus Bolanderi (Gray) Greene, and Salvia
columbariae Benth. The species name is derived from a Greek
word meaning most beautiful.

This plant is related to Streptanthus hispidus Gray, known only
from Mt. Diablo, from which it is readily separated by the short,
terete, curved pods with small, rounded, wingless seeds. Strept-
anthus insignis Jepson, known from the inner South Coast Ranges
of western Fresno, San Benito and eastern Monterey counties, is
also related to Streptanthus callistus by reason of the similarity in
the terminal, sterile flowers. These three related species may be
distinguished by the following characters:

Siliques flattened, straight, erect, 5.0—7.0 cm. long; seeds
winged, 0.2 cm. long.

Plants densely hispid, low, compact, 5.0-30.0
cm. high; leaves spatulate-oblanceolate,
saliently lobed, reddish or purple below;
blade of petal light purple, calyx densely
hispid, sterile flowers either very rarely
present or slightly developed, not forming
a terminal color spot; stigma entire ..... Streptanthus hispidus

Plants sparsely hispid, slender, 10.0—-45.0 cm.
high; leaves deeply lobed, almost pinnati-
fid; blade of petal white, with a black-
purple midvein, calyx purple, sparsely
hispid, terminal cluster of sterile flowers
always present, black purple or garnet,
forming a prominent terminal color spot;

stigma slightly two: lobed ....0). 7.2.5.4 Streptanthus insignis
Siliques terete, incurved, erect, 1.5-2.0 cm. long; seeds
not flattened, wingless, 0.1 cm. long .............. Streptanthus callistus

University of California, Berkeley,
March 15, 1938.

1938 ] SHARSMITH: COREOPSIS 209

THE NATIVE CALIFORNIAN SPECIES OF THE
GENUS COREOPSIS L.

Heten K. SuHarsmMitH

For the past three years the writer has been engaged in a

floristic survey of the Mount Hamilton Range, a northern unit of

the inner South Coast Ranges of California. The interior and
eastern regions of the Mount Hamilton Range are comparatively
isolated, and in the past they have received little botanical explo-
ration. Recent field work has resulted, therefore, in the accumu-
lation of information on several new or little known species of
Californian plants. This paper is an outgrowth of field acquain-
tance with the four species of Coreopsis which grow in the Mount
Hamilton Range. |

From the time of De Candolle to recent years, the Califor-
nian species of Coreopsis have had a varied generic history in-
volving Leptosyne DC., Agarista DC., Tuckermannia Nutt., and
Pugiopappus Gray. In the Synoptical Flora of North America
(17: 299-301. 1884), Gray included these four genera, along
with Coreocarpus Benth. and Acoma Benth., in an amplified ver-
sion of Leptosyne, limiting the genus Coreopsis to the eastern side
of the continent. This was the accepted interpretation of many

subsequent botanists. Harvey M. Hall (Univ. Calif. Publ.

Bot. 3: 189-143. 1907) followed Bentham and Hooker (Genera

Plantarum 2: 385. 1878) and O. Hoffmann (in Engler and
Prantl, Natirlichen Pflanzenfamilien 4°: 243. 1894) in uniting
Leptosyne with Coreopsis. In “A redisposition of the species
heretofore referred to Leptosyne” (Proc. Am. Acad. 49: 335-346.
1913), S. F. Blake amply substantiated this viewpoint, but re-
moved Coreocarpus, a Sonoran genus of three species, from the
aggregate genus Coreopsis. His paper gives a complete presen-
tation of the generic and intrageneric relationships involved.
According to his conception the native Californian species of the
genus Coreopsis belong to the subgenus Leptosyne (DC.) Blake.
This subgenus also embraces six Mexican species and reaches
south as far as Guatemala.

The eight Californian species of Coreopsis, including six an-
nuals and two perennials, are almost entirely restricted to the
botanical province of California. They are most abundant in
the cismontane and desert areas of southern California, and only
C. Stillmanit reaches north of central California. Two of the
eight species, C. maritima and C. californica, transgress the politi-
cal boundaries of the state, but neither is found any considerable

distance beyond. The accompanying map (text fig. 1) repre-

sents the distribution of the genus in California.
Field observations have indicated that fresh material is nec-
essary for adequate taxonomic treatment of the annual species

210 MADRONO [Vol. 4

C. BIGELOVI)
C.HAMILTONII
C.CALLIOPSIDEA
C.CALIFORNICA
C. DOUGLASI|I
C.STILLMANII
C.MARITIMA

: C.GIGANTEA

opp BeOXxXoO

°o
7 o%o
ae ‘ oe ene® 0° o 2 ;
Pa meas y 0085 2 pe Pad 2,
Q Oy oe, ARIZONA
ee
Oo tare 4 e © ee
e
CALIFORNIA Beis ori ors
@ - yee
‘hn

e
BAJA CALIFORNIA

) e

Fig. 1. Distribution of native Californian species of Coreopsis.

of Coreopsis found in California. Many of the distinguishing
characters, particularly those of the leaves and of the involucre,
are lost in the pressed specimens. All six annual species were
studied from fresh material, either from that gathered in the
field or from garden cultures. In addition heads preserved in
formalin were obtained for two of the southern Californian
species. |

In addition to field observations and study of fresh material,
herbarium specimens were borrowed from the following Cali-
fornian institutions, and appreciation is expressed to the cura- .
tors of these herbaria: California Academy of Sciences, San
Francisco (CA), Dudley Herbarium, Stanford University (SU),
Pomona College, Claremont (P), Santa Barbara Museum of

1938 | SHARSMITH: COREOPSIS 211

Natural History (SBM), University of California (UC), Univer-
sity of California at Los Angeles (UCLA), Vegetation Type Map
Herbarium, California Forest and Range Experiment Station,
Berkeley (VTM).

Acknowledgment is due to the following persons for the
generous responses made to the requests of the writer: Dr. G.
Ledyard Stebbins, Jr., and Dr. Herbert L. Mason of the Univer-
sity of California, Berkeley; Dr. Carl Epling, University of Cali-
fornia at Los Angeles; Mr. Frank W. Peirson, Altadena, Cali-
fornia; Mr. Maunsell Van Rensselaer, Blaksley Botanic Garden,
Santa Barbara, California; Sir Arthur W. Hill, Director, Royal
Botanic Gardens, Kew, England.

Key To THE NATIVE CALIFORNIAN SPECIES OF COREOPSIS

Annuals; stems slender, 5-60 cm. tall; cismontane and desert species.
Achenes dimorphic; disc achenes ciliate, pappus paleae 2, ray achenes gla-
brous, epappose. Section Pugiopappus (Gray) Blake.

Outer series of involucral bracts linear, obtuse, shorter to longer than
the ovate inner series; leaves mostly basal and scapes naked.

Ligules horizontal in anthesis; receptacular bracts falling attached
to disc achenes; palea of disc achenes mostly 2 mm. long.

1. C. Bigelovit.

Ligules strongly reflexed in anthesis; receptacular bracts falling
separately from disc achenes; palea of disc achenes 1 mm. long.

2. C. hamiltonia.

Outer series of involucral bracts broadly ovate, acute, shorter than

the oblong-ovate inner series; lower 3 to 3 of stems leafy, leaves

only slightly clustered at base; palea of disc achenes mostly 4 mm.

OTN eee ie Pe ee ec Oe ia ek SO ne ae bes 3. C.calliopsidea.

Achenes monomorphic, non-ciliate; pappus reduced to a cupule. Section

Euleptosyne (Gray) Blake.

Leaves linear to filiform, fleshy, 1 mm. wide, entire or with 1-2 linear
pinnae, terminal lobe not broader; annulus of disc corollas
bearded; outer involucral bracts narrowly lanceolate, gibbous
at base.

Leaves filiform, terete, only slightly flattened above, erect; achenes
dull, tan to light brown or reddish, roughened with clavellate
or capitate hairs on body and wing, and with a central corky
ridge, corky wing irregularly thickened...... 4. C. californica.

Leaves linear, grooved above, rounded beneath, spreading;
achenes shining, body dark brown, smooth, glabrous or with
a few coarse hairs or callous papillae, corky wing light tan, thin.

5. C. Douglasii.

Leaves flat, only slightly fleshy, pinnate or bipinnate into spatulate

lobes 1-3 mm. broad, terminal lobe usually broadest; annulus of

disc corollas mostly glabrous; outer involucral bracts linear to

linear-spatulate, not gibbous at base............ 6. C. Stillmanii.

Perennials; stems stout, 3-30 dm. tall; coastal and insular. Section Tucker-
mannia (Nutt.) Blake.

Heads few (2-4) at ends of branches, on naked peduncles 15-50 cm. long;

coastal San Diego County to northern Baja California and adjacent

TS ING Sareea Tay Li oe ce te ne Maat AB tc soit e-dtyh, A aes 7. C. maritima.

Heads numerous and cymosely clustered at ends of branches, on leafy
peduncles 6-20 cm. long; coastal southern California and adjacent
islands from San Luis Obispo County to Los Angeles County.

8. C. gigantea.

212 MADRONO [Vol. 4

1. Coreopsis Bigetovu (Gray) Hall, Univ. Calif. Publ. Bot. 3:
141. 1907. Pugiopappus Bigelovii Gray, Pac. R. Rep. 4: 104.
1857. P. Breweri Gray, Proc. Am. Acad. 7: 660. 1878. Lepto-
syne Bigelovii Gray, Syn. Fl. 1, pt. 2: 800. 1884.

Erect, essentially glabrous annual with several (1-30, mostly
5-10) scapose stems from a taproot: scapes yellowish green to
stramineous, somewhat leafy below and occasionally branching
near base, sometimes with 1 or 2 appressed bracts above the
leaves, monocephalous, 10-60 (mostly 25-30) cm. tall, usually
stout: leaves 4-12 cm. long, alternate, basally clustered, only
slightly fleshy, flattened, lightly grooved above, rounded below,
somewhat glaucous, spreading; blade ovate, 2-8 cm. long, once
or twice pinnate into linear, obtuse, often callous tipped lobes
5-30 mm. long and 1-2 mm. wide, rachis as wide as lobes; peti-
ole 1-5 mm. long, 1-2 mm. wide, channelled, expanded at base,
sometimes irregularly margined with minute glandular capitate
hairs: heads erect, 1-1.5 em. high, 2—4.5 ecm. wide: involucre
glabrous, cylindric or slightly barrel shaped, with sharply trun-
cate base; bracts of outer series 4-7, linear, 5-12 mm. long
(often longer than inner bracts), dark green, glaucous to shin-
ing, slightly fleshy, mostly loosely spreading from base; bracts
of inner series 6—8, ovate, 6-10 mm. long, 83-6 mm. wide; acute
with apical tuft of hyaline hairs, greenish yellow, shining, many
nerved, scarious margined, erect or sometimes with tips spread-
ing, reflexed in fruit: ray florets 5-10, mostly 8, pistillate, fer-
tile; ligule golden yellow, obovate, 5-25 (mostly 15) mm. long,
3-12 (mostly 8) mm. wide, with truncate and erose apex, spread-
ing horizontally; tube 2 mm. long with scattered glandular capi-
tate hairs; style branches linear, obtuse: dise florets 20—50, per-
fect, golden yellow, all fertile, or the central sometimes sterile;
tube 2 mm. long; annulus glandular pubescent; throat expanded,
1.5 mm. long; corolla lobes triangular ovate, acute, glandular
papillate, spreading; anther tips ovate cordate; style tips tri-
angular acute, glandular papillate; pollen grains spherical, with
short spines: receptacular bracts lanceolate to oblanceolate, 4-10
(mostly 5-8) mm. long, 1 mm. broad, subacute to acute, hyaline,.
slightly callous thickened on midrib, exceeding body of disc
achenes and often equalling the pappus palea, closely adhering
to back of and persistently attached to base of disc achenes: ray
achenes obcompressed, oblong to obovate, 3-5 mm. long, 2 mm.
wide, glabrous, epappose; body brown or mottled with tan, often
almost covered with tan, corky, microscopically foveolate rough-
enings, flat or somewhat ridged on inner face, rounded on back;
marginal callous wing narrow, smooth or corky roughened simi-
larly to but more densely than body: dise achenes obcompressed,
oblong to oblanceolate, 4-6 mm. long, 1-1.5 mm. wide; body
dark brown, sometimes mottled with tan, shining, microscopi-
cally latticed, truncate at base with an evident callous; back

1938 | SHARSMITH: COREOPSIS 213

completely glabrous, covered with the persistent receptacular
bract; inner face flat or with slight central ridge, glabrous ex-
cept for a central row of more or less prominent cilia similar to
but shorter than marginal cilia; marginal cilia silky, white, up-
wardly directed, two-celled, microscopically bidentate at apex,
1-1.5 mm. long; pappus palea of two, persistent, lanceolate, tri-
quetrous, acuminate, hyaline, denticulate membranes 2 (or 3)
mm. long.

Range. Desert and cismontane areas of southern California
and the southern Sierra Nevada of California, altitude 150-1500
m.; from San Diego County north to Tulare County, west to
Santa Barbara County, and north to southern Monterey County.

Type. “On the Mohave Creek, in the desert east [west] of
Colorado,’ March, 1854, J. M. Bigelow.

Specimens examined. San Diego County: Colorado Desert,
W.G. Wright (UC). Riverside County: near Banning, April 238,
1922, M. F. Spencer 2040 (P). San Bernardino County: Bar-
stow, May 1, 1922, M. F. Spencer 1954 (P); 7 miles east of Dag-
gett, April 6, 1924, P. A. Munz and D. Keck 7842 (P); Mescal
Creek, April 29, 1902, H. M. Hall 3027 (UC); Cajon Pass, May,
1905, H. M. Hall 6219 (UC); Trona road, May 8, 1937, An-
netta Carter 1305 (UC). Los Angeles County: Mandeville Can-
yon, March, 1929, I. W. Clokey and B. Templeton 4520 (P);
Saugus, April 23, 1903, G. B. Grant 5424 (P, UC); Antelope
Valley, May 1-8, 1902, H. M. Hall 3074 (UC); 5 miles south of
Muroc, April 1, 1932, H. L. Mason 6882 (UC); between Rock
Creek and Little Rock Creek, April 27, 1926, H. L. Mason 3046
(UC). Kern County: between Mohave and Lancaster, April 20,
1930, H. L. Mason 5567 (UC); Greenhorn Range, June 2-10,
1904, H. M. Hall and H. D. Babcock 5078 (UC); Kern Canyon,
April 12, 1905, A. A. Heller 7662 (UC); vicinity of old Fort
Tejon, June 16-19, 1905, H. M. Hall 6307 (UC). Inyo County:
April 17, 1891, T. S. Brandegee (UC); 10 miles sw. of Shoshone,
April 17, 1982, C. L. Hitchcock 12340 (P, UC); Argus Mountains,
April 18, 1930, Harold Bailey and W. Robison (P, UCLA, UC).
Tulare County: near Oak Grove, April 23, 1925, R. Bacigalupi
1209 (P); Kaweah River Basin, April, 1901, Ralph Hopping 38
(UC); Tule River, March 22, 1925, P. A. Munz 91386 (P). Ven-
tura County: Pine Creek near Sespe, March 24, 1917, B. W. Ever-
man (CA). Santa Barbara County: Figueroa Mountain, May
31, 1929, Ralph Hoffmann (SBM); Dutch Flat, San Rafael Moun-
tains, June 8, 1929, Ralph Hoffmann (SBM); trail to Zaca Peak,
June 19-80, 1906, Alice Eastwood 585 (CA). Monterey County:
north of Parkfield, March 24, 1925, P. A. Munz 9186 (P).

Coreopsis Bigelovit was described by Gray as having the annu-
lus of the disc corollas beardless. A similar interpretation was
given by Hall and also by Jepson (Man. Fl. Pl. Calif. 1084,
1085. 1925) who uses the beardless annulus as one of the char-

214 MADRONO [Vol. 4

acters distinguishing C. Bigelovii from C. calliopsidea. Blake,
after an examination of the type of C. Bigelovii, states (op. cit.,
p. 341): “The annulus of the disk-flowers, in the types and other
specimens examined, is very distinctly bearded, not glabrous as
originally described and as repeated in the Synoptic Flora and
by Hall.” The specimens cited above verify this observation of
Blake’s, though in some of the specimens the ring of hairs is not
complete.

2. Coreopsis hamiltonii (Elmer) comb. nov. Leptosyne
hamilton Elmer, Bot. Gaz. 41: 3828. 1906.

Erect, essentially glabrous annual with several (1-80, mostly
8-10) scapose stems from a taproot: cotyledons linear, 1.5 cm.
long, 1 mm. wide, flattened, spreading: scapes reddish, somewhat
leafy below and occasionally branching near base, sometimes with
a small, appressed bract above the leaves, monocephalous, 6—25
(mostly 10-15) em. tall: leaves 1-5 cm. long, alternate, basally
clustered, fleshy, flattened and lightly grooved above, rounded be-
low, glaucous, spreading; blade triangular ovate, 0.5—2 cm. long,
often as wide, twice pinnate into linear, obtuse and sometimes
callous tipped lobes 1 mm. wide; petiole 1-3 cm. long, 2 mm.
wide, strongly grooved above, expanded at base: heads erect,
7-10 mm. high, 10-20 mm. wide: involucre glabrous, cylindric
or somewhat barrel shaped, with sharply truncate base; bracts
of outer series 4-7, linear, 3-6 mm. long, always shorter than
inner series, obtuse, dark green occasionally splotched with red,
glaucous to shining, slightly fleshy, appressed to inner bracts
and spreading only near tips; bracts of inner series 6—8, broadly
lanceolate or narrowly ovate, 5-8 mm. long, 2—3 mm. wide, acute
with apical tuft of hyaline hairs, greenish yellow, shining, thin,
many nerved, narrowly scarious margined, appressed or some-
times with tips spreading, reflexed in fruit: ray florets 5-8,
pistillate, fertile; ligule golden yellow, oblong to obovate, 3-8
mm. long, 2-5 mm. wide, obscurely 3-toothed at the truncate
apex, strongly reflexed; tube 1.5—2 mm. long with incomplete
ring of short, yellowish, glandular capitate hairs near apex;
style branches linear, obtuse: dise florets 20-30, perfect, golden
yellow, central ones sterile; tube 1-2 mm. long, annulus glandu-
lar pubescent; throat expanded, 1.5-2 mm. long; corolla lobes
triangular ovate, acute, glandular papillate, spreading; anther
tips ovate cordate; style tips triangular acute, glandular papil-
late; pollen grains spherical, with short spines: receptacular
bracts linear, 5-6 mm. long, 1 mm. broad, obtuse, hyaline, falling
separately from dise achenes: ray achenes obcompressed, menis-
coid, obovate, 5 mm. long, 8 mm. wide, dark brown or tan
splotched with brown, smooth, glabrous and shining, rounded on
back, flat on inner face with slight central keel epappose; marginal
wing thin, smooth, tan marbled with brown: disc achenes obcom-
pressed, narrowly obovoid, 5-6 mm. long, 1.5-2 mm. broad,

1938 ] SHARSMITH: COREOPSIS 215

smooth, tan marbled with brown, truncate at base with slight
callous, rounded on both sides, the inner face with slight central
ridge or keel, both faces covered (sometimes sparsely so) with
persistent, upwardly directed, silky, white, two-celled cilia 1
mm. long, which are microscopically bidentate at apex and often
swollen at base, the marginal cilia thicker, more spreading, and
up to 1.5 mm. long; pappus palea of two, persistent, obovoid,
triquetrous, triangular acute, hyaline, denticulate membranes 1
mm. long.

Range. Exposed, dry rocky slopes of Santa Clara County
in the Mount Hamilton Range of the inner South Coast Ranges
of California, altitude 600-1295 meters.

Type. Mount Hamilton, Santa Clara County, California,
April, 1900, 4. D. E. Elmer 2328. The collector comments as
follows: “It was in fine flower and fruit, and grew in dry gravelly
soil on a steep slope a few hundred yards below the observatory.
Since then I have failed to find it either in this same place or
elsewhere.”

Specimens examined. Santa Clara County: Mount Hamil-
ton, April 29, 1923, Alice Eastwood 11671, topotype (CA) ; Coper-
nicus Peak, highest point of Mount Hamilton, altitude 4250 feet,
April 15, 19384, H. K. Sharsmith 914, topotype, representative of
the species (UC); April 28, 19385, H. K. Sharsmith 1839, topotype
(UC); Mount Hamilton-Livermore road, April 28, 1925, Alice
Eastwood 12468 (CA); eight miles from Mount Hamilton on road
to San Antonio Valley, April 6, 1930, J. T. Howell 4665 (CA),
April 6, 1930, FE. K. Crum 602 (UC); San Antonio Valley, March
23,1982, H. HE. Wieser (SU); Arroyo Bayo, March 80, 1935, H. K.
Sharsmith 1709, April 2, 1936, 3489, April 25, 1986, 8628 (UC).

Coreopsis hamiltonit has been overlooked as a specific entity
ever since Elmer’s description of it in 1906. The only recent
reference occurs in Blake’s paper (op. cit., p. 841), in which the
original combination (Leptosyne hamiltonii Elmer) is given as a
synonym under C. Bigelovii; no specimens are cited and no local-
ity is mentioned. Elmer’s original description of C. hamiltonii
is detailed in content and presents, in general, an adequate pic-
ture of the species. It is in error, however, regarding the
- achenes, which are described as “ciliate on the edges, pubescent
on the sides, brown and glabrous when mature: pappus of two
caducous hyaline finely ciliate membranes.’ Actually, the
achenes are dimorphic, those of the ray differing strongly from
those of the disc.

3. COREOPSIS CALLIOPSIDEA (DC.) Gray, Bot. Mex. Bound. 90.
1859. Agarista calliopsidea DC. Prod. 5: 569. 18386. Pugio-
pappus calliopsidea Gray, Proc. Am. Acad. 8: 660. 1873. Lepto-
syne calliopsidea Gray, Syn. Fl. 1, pt. 2: 800. 1884. JL. calli-
opsidea var. nana Gray, loc. cit.

Erect, essentially glabrous, usually rather stout annual with

216 MADRONO [Vol. 4

several (1-30 or more, mostly 4-6) stems from a taproot: stems
yellowish green to stramineous, usually leafy to the middle or
even above, sometimes almost naked, sometimes branching near
the base, occasionally branching above, monocephalous, 8-55
(mostly 20-80) cm. tall: leaves 4-8 em. long near base, reduced
above, alternate, somewhat clustered at base or quite scattered
on stems, only slightly fleshy, lightly grooved above, rounded
beneath, somewhat glaucous, spreading; blade ovate, 1—5 em.
long, once or twice pinnate into linear, obtuse, divergent lobes
0.5-2 mm. wide and often callous tipped, rachis as wide as lobes
or up to 5 mm. wide, upper leaves sometimes simple; petiole 1-5
cm. long, 0.5-5 mm. wide, channelled, expanded at base: heads
erect, showy, 1-2 cm. high, 2-9 cm. wide: involucre campanu-
late, united at base into a disc 3-10 mm. across, this disc glabrous
or with a few, scattered glandular hairs; bracts of outer series 5
(occasionally 4 or 6), deltoid ovate with a broad base, 3-8 mm.
long, almost as wide to wider, acute and sometimes callous
tipped, green, glaucous, flat, not fleshy, margined occasionally
with short glandular capitate hairs, somewhat spreading; bracts
of inner series mostly 8 (occasionally 6 or 7), ovate with a broad
base, 8-10 mm. long, exceeding the outer series, acute with a
minute, apical tuft of hyaline hairs, greenish yellow, shining,
thin, narrowly scarious margined, many nerved, somewhat
spreading: ray florets mostly 8 (occasionally 6 or 7), pistillate,
fertile or often sterile; ligule golden yellow, obovate, 10-35 mm.
long, 5-18 mm. broad, with truncate erose apex, spreading hori-
zontally; tube 8 mm. long, essentially glabrous; style branches
linear, obtuse: dise florets 15—50, perfect, golden yellow, usually
all fertile; tube 2 mm. long, with scattered glandular hairs;
annulus heavily bearded with glandular hairs; throat funnelform,
2 mm. long; corolla lobes lanceolate, 1 mm. long, acute, glandu-
lar papillate, spreading; anther tips lanceolate cordate; style
tips triangular acuminate, glandular papillate; pollen grains
spherical, with short spines; receptacular bracts lanceolate to
oblanceolate, 6-7 mm. long, 1-1.5 mm. wide, acuminate to apicu-_
late, hyaline, the midrib callous thickened, scale equalling or
searcely exceeding body of disc achene, closely adhering to its
dorsal surface and persistently attached to its base: ray achenes
(if developed) obcompressed, oval, 5-6 mm. long, 3-4 mm. wide,
glabrous, epappose; body tan or brown, smooth or sometimes
almost covered with a rough, tan callous, flat or somewhat
ridged on inner face, rounded on back; marginal wing smooth,
stramineous, flat: disc achenes obcompressed, linear to oblance-
olate, 6-7 mm. long, 1.5 mm. wide; body dark brown, shining,
microscopically latticed, rounded at base with an evident callous;
inner face flat, covered with upwardly directed, silky, white or
tawny, two-celled cilia 1 mm. long and microscopically bidentate

1938 | SHARSMITH: COREOPSIS 217

at apex; back somewhat rounded, completely glabrous, covered
with the persistent receptacular bract; marginal cilia thicker, 2—3
mm. long; pappus palea of two, persistent, lanceolate, triquetrous,
acuminate, hyaline, denticulate membranes 2.5—5 (mostly 4) mm.
long.

Range. Western portion of the Mohave Desert in San Ber-
nardino County and Kern County; west to eastern San Luis
Obispo County; north on extreme eastern side of inner South
Coast Ranges to Corral Hollow, Mount Hamilton Range, Ala-
meda County, altitude 210-1070 meters.

Type. “In California legit cl. Douglas.”

Specimens examined. San Bernardino County: Mohave Des-
ert, May, 1882, C. G. Pringle, isotype of C. calliopsidea var. nana
Gray (UC); Crutts Post Office, May 14, 1922, 7. M. Johnson 6475
GUC); Fremont’s Peak, May 6, 1906, H. M. Hall and.H. P.
Chandler 6868, 6869 (UC) ; near Palmdale, March 28, 1931, F. A.
MacFadden 2574 (UC). Kern County: Mohave Station, June 38,
1915, S. B. Parish 9771 (UC); Randsburg, April 5, 1927, Craig,
Newsom, Hilend 129 (P); Bakersfield, April 4, 1893, Alice East-
wood (UC). Los Angeles County: 3 miles west of Muroc, Mo-
have Desert, April 2, 19382, H. L. Mason 6895 (UC). Santa Bar-
bara County: Cuyama, White Hills, May 2, 1896, Alice Eastwood
(UC). San Luis Obispo County: Carrizo Plains, April 1, 19384,
E. Armstrong 1102 (VTM). Fresno County: Alcalde, April,
1891, T. S. Brandegee (UC); Panoche Creek, April 1, 1982, C. H.
Quibell 2799a (P). Stanislaus County: Arroyo del Puerto, Mount
Hamilton Range, altitude 730 feet, March 29, 1935, H. K. Shar-
smith 15938 (UC). Alameda County: Tesla, April 11, 1935, J. T.
Howell 12598 (CA); Corral Hollow, April 21, 1935, A. M. Carter
mane (UC). |

Coreopsis calliopsidea is the only one of the annual species here
treated in which the stems are always somewhat leafy. Basal
clustering of the leaves may be quite absent, and the stems may
be clothed with scattered leaves for their greater length, or the
upper half or two-thirds of the stems may be essentially naked.
Coreopsis calliopsidea var. nana Gray was based on small plants in
which the leaves are somewhat basally crowded, and the stems
more scapose than usual. In Hall’s treatment (op. cit., p. 141)
the outer disc achenes are described as being like those of the
ray, oval, flat, and glabrous, the central disc achenes as being
narrow with long cilia. The dimorphism, however, is sharply
defined between achenes of ray and disc, in this species as in the
other members of the section Pugiopappus. Only the ray achenes
are glabrous, and all of the disc achenes are ciliate.

4. Coreopsis californica (Nutt.) comb. nov. Leptosyne cali-
fornica Nutt. Trans. Am. Phil. Soc. n. ser. 7: 3638. 1841. L. New-
berryt Gray, Proc. Am. Acad. 7: 358. 1868. L. Douglasii of

218 MADRONO [Vol. 4

Gray and other authors, not L. Douglasii DC. Coreopsis Douglasii
Hall, Univ. Calif. Publ. Bot. 3: 140. 1907 (misapplied).

Erect, essentially glabrous annual with several (1—20, mostly
about 5) scapose stems from a slender taproot: cotyledons linear
filiform, 83-4 cm. long, obtuse with a conspicuous red callous tip,
fleshy, terete or slightly flattened above, erect: scapes slightly
reddish, somewhat leafy and occasionally branching near base,
often with a small, appressed bract above the leaves, mono-
cephalous, 5-45 (often 20-25) cm. tall: leaves basally clustered,
erect, linear filiform, 2-10 (or 15) cm. long, 0.5 mm. wide, ob-
tuse with a conspicuous red callous tip, fleshy, terete or slightly
flattened above, mostly entire, occasionally with 1—2 small, linear
pinnae, glabrous except for an occasional, short, glandular capi-
tate hair: heads erect and solitary, 6-10 mm. high, 10-35 mm.
broad: involucre barrel shaped with rounded base; bracts of
outer series 2—7, pale green, sometimes blotched with red, promi-
nently bearded at gibbous base with vellow or red glandular
papillae, slightly fleshy, narrowly lanceolate, 4-7 mm. long, 1
mm. wide, obtuse with a red callous tip, spreading; bracts of
inner series 5—8 (or 10), broadly lanceolate to narrowly obovate,
6-10 mm. long, 2—4 mm. wide, acute with tuft of apical hyaline
hairs, yellow green often blotched with red, thin, many nerved,
narrowly scarious margined, constricted at middle with upper
half spreading in anthesis, connivent in early fruit, spreading in
age: ray florets 5-12, pistillate, fertile; ligule yellow, sometimes
paler near apex, narrowly to broadly obovoid, 5-15 mm. long,
3-8 mm. wide, strongly 3-toothed at apex, spreading horizon-
tally; tube 1-1.5 mm. long with apical ring of brownish or yel-
lowish glandular papillae; style branches linear, obtuse: disc
florets 10-30, perfect, yellow, central ones often sterile; tube
1-2 mm. long; annulus glandular pubescent with yellowish or
brownish hairs; throat expanded, 2 mm. long; corolla lobes
broadly ovate, acute, spreading; anther tips ovate cordate; style
tips triangular, attenuate, glandular papillate; pollen grains
spherical, with short spines; receptacular bracts linear to spatu-
ulate, 4-55 mm. long, 1-1.5 mm. broad, obtuse, hyaline, lightly
nerved, not falling with disc achenes and often more tardily
deciduous: achenes monomorphic, obcompressed, obovate, 2.5—5
mm. long, dull and microscopically papillate, roughened with
short, hyaline, two-celled, clavellate or capitate hairs on wing
and body, rusty-tan to light brown, tardily deciduous; pappus
reduced to a cupule; marginal wing of achene colored like body
or slightly lighter, strongly and irregularly corky thickened and
spongy, microscopically foveolate, with marginal row of hyaline,
two-celled, clavellate or capitate hairs and often with row of 1 to
several red spots on inner edge; body of achene rounded on back,
inner face with a well developed, central corky ridge; central

1938 ] SHARSMITH: COREOPSIS Zio

sterile achenes varying to thin, smooth, somewhat shining, only
sparsely hairy, usually with clavellate hairs forming an obvious
marginal fringe on the wing, light tan or stramineous, often with
conspicuous and numerous red spots along inner edge of wing.
Range. Cismontane southern California, Mohave Desert and
Colorado Desert, from southeastern Santa Barbara County to
San Diego County, altitude 30-600 meters; south to northwest
Lower California, east to southern Arizona.
Type. ‘Near San Diego. Upper California,” Nuttall.
Specimens examined. Catirornia. Santa Barbara County:
Le Cumbre Club, R. C. and J. Robbins in 1921 (SBM). Los
Angeles County: Little Rock, April 26, 1926, H. L. Mason 3003
(UC); Lancaster, April 1, 1932, H. L. Mason 6872 (UC); Pasa-
dena, May 2, 1882, Marcus E. Jones 3361 pro parte, 3373 (P);
Claremont, May 20, 1909, L. Abrams 53825 (SU); Cobal Canyon,
March 16, 1925, P. A. Munz 9357 (P); near Santa Ana Canyon,
March 10, 1926, M. E. Jones (P); Altadena, April, 1902, G. B.
Grant 681 (UC); Verdugo Hills, April 6, 1901, L. Abrams 1387
(SU). Orange County: Santa Ana Mountains, April, 1904, H.
D. Geis 732 (SU). Kern County: El Paso Range, May 1, 1927,
L. R. Abrams 11915 (P); 10 miles east of Mohave, April 2, 1932,
H. L. Mason 6906 (UC); between Rosamund and Mohave, April
30, 1927, L. R. Abrams 11788 (P, SU); between Coso Hot
Springs and Coso Junction, April 30, 1928, R. S. Ferris 7466 (P).
San Bernardino County: north of Box S Ranch, April 21, 1932,
P. A. Munz 12415 (P, UC); between Warren’s Well and Coyote
Well, April 18, 1930, J. A. Ewan 3057 (UC); 20 miles northwest
of Barstow, May 5, 1906, H. M. Hail and H. P. Chandler 6850
(UC) ; west of 29 Palms, April 10, 1935, P. A. Munz 138774 (P);
summit Cajon Pass, April 12, 1919, S. B. Parish 19237 (UC).
Riverside County: Riverside, April 4, 1902, H. M. Hall 2960
(UC); se. of White Tanks, April 9, 1932, P. A. Munz and C. L.
Hitchcock 12224 (P, UC). Imperial County: Coyote Canyon,
April, 1902, H. M. Hall 2859 (UC, SU). San Diego County: San
Diego, April 21, 1908, H. M. Hall 3860 (UC); Del Mar, March
28, 1894, T. S. Brandegee (UC); Point Loma, April 6, 1913, Alice
Eastwood 2533 (CA); Jacumba Springs, April 11-16, 1924, W. W.
Eggleston 19748 (P). Arizona. Arnett Canyon, Pinal County,
March 19, 1932, J. W. Gillespie 5420 (UC); Apache Trail, March
6, 1929, Mrs. C. W. McKelvey (CA). Mexico. Baja California:
Northwest corner Baja California, April, 1903, H. M. Hall 3973
(UC) ; San Quintin, February 15, 1935, C. Epling and W. Robison
(UCLA) ; Todos Santos Bay, April 10, 1882, M. E. Jones (SU).
From the time of Gray to the present, the southern Cali-
fornian species described above as C. californica has been known,
first, as Leptosyne Douglasii, and later, as Coreopsis Douglasii.
Soon after beginning the present study it became evident that
there are two species among the plants recognized as C. Douglasii,

220 MADRONO [Vol. 4

the most outstanding differences being in the fruits. Through
the courtesy of Sir Arthur Hill, two fruits from the type of
Leptosyne Douglasii DC. were sent for comparison from the Royal
Botanic Gardens at Kew. Though somewhat immature, these
fruits belong unquestionably, not to the widely spread and well
known southern California unit under consideration, but to the
relatively restricted and hitherto unrecognized unit of the inner
South Coast Ranges. The name C. Douglasii must be transferred,
then, from its old and well-known association with the southern
Californian species, to the South Coast Range species. It follows
that Leptosyne californica must be reestablished as the type of the
southern Californian species.

According to Nuttall’s description of Leptosyne californica, the
ray achenes are imperfectly developed and almost smooth, but
the disc achenes are scabrous with short, glandular hairs, and
have, as well, thick, spongy margins and often a similarly en-
larged center. Leptosyne Newberryi Gray, described from mate-
rial collected by Newberry at Sitgreaves Pass, Arizona, and by
Palmer at Camp Grant, Arizona, represents a phase in which the
achenes are only sparsely clad with clavellate hairs. These
minor achenal differences give evidence of the variations to be
expected in the achenes of C. californica. If only the extremes
are considered, the differences are striking. The variation is not,
however, one of regional significance, but seems, rather, to be
related to fertility within the individual head. The central disc
florets of this species are more or less sterile and vary from the
fertile ray and outer dise achenes in the manner indicated in the
description of the species. Both extremes may be present in the
same head, or the more fertile type may be quite lacking.

When there is taken into consideration the wide range of
climatic and edaphic factors to which C. californica is exposed, the
size variation which occurs within the species is not surprising.
The species is sufficiently plastic to adapt itself to habitat condi-
tions as diverse as the Mohave Desert and the seacoast mesas of
San Diego County. The maritime form often has the basal leaves
thickly clustered, and the scapes stout and rather low with large
and showy heads.

5. Coreopsis Doverasu (DC.) Hall, Univ. Calif. Publ. Bot. 3:
140. 1907 (as to name but not as to description). Leptosyne
Douglasii DC. Prod. 5: 581. 1886. C. Stillmanii var. Jonesu
Sherff, Bot. Gaz. 97: 605. 19386.

Erect, essentially glabrous, glaucous annual with several (1—
12, mostly 2—5) stems from a slender taproot: cotyledons linear,
2—5 cm. long, obtuse with an inconspicuous red callous tip, fleshy,
rounded beneath, flattened and lightly grooved above, spread-
ing: stems reddish, scapose, somewhat leafy and occasionally
branching near base, sometimes with a small, appressed bract

1938 ] SHARSMITH: COREOPSIS 221

above basal leaves, monocephalous, 5-25 (mostly 15) cm. tall,
glabrous or with an occasional, short, glandular capitate hair:
leaves alternate, basally clustered, spreading when young, sub-
erect to erect with age, linear, 2-8 cm. long, 1 mm. wide, obtuse
with an inconspicuous red callous tip, fleshy, rounded beneath,
flattened and lightly grooved above, entire or with 1-2 linear
pinnae 3-10 mm. long, glabrous except for lateral rows of widely
scattered, short, glandular-capitate hairs: heads often nodding in
bud, erect in flower, 6-10 mm. high, 10-25 mm. broad; involucre
barrel shaped with rounded base; bracts of outer series 2—7, pale
green, inconspicuously bearded with short, yellowish glandular
papillae at gibbous base, slightly fleshy, narrowly lanceolate, 4—7
mm. long, 1 mm. wide, obtuse, spreading; bracts of inner series
5-8, obovate, 6-10 mm. long, 3—4 mm. wide, acute with apical
tuft of hyaline hairs, pale green to yellowish or reddish green,
thin, many nerved, narrowly scarious margined, constricted at
middle with upper half spreading in anthesis, connivent in early
fruit, spreading in mature fruiting heads: ray florets 5-8, pistil-
late, fertile; ligule golden yellow, sometimes paler near apex,
oval, 5-8 mm. long, 3-5 mm. wide, weakly 3-toothed at apex,
spreading horizontally, tube 1.5-2 mm. long with hairy ring of
glandular hairs at apex; style branches linear, obtuse: dise florets
10-30, perfect, golden yellow, central ones often sterile; tube
1.5—2 mm. long; annulus glandular pubescent; throat expanded,
2mm. long; corolla lobes broadly ovate, acute, glandular papil-
late, spreading; anther tips ovate cordate; style tips triangular,
attenuate: pollen grains spherical, with short spines: recep-
tacular bracts linear, 4-5 mm. long, 1-2 mm. wide, obtuse, hya-
line, lightly nerved, falling separately from disc achenes: achenes
monomorphic, obcompressed, obovate, meniscoid, 2.5-5 mm.
long, essentially glabrous, smooth, somewhat tardily deciduous;
pappus reduced to a cupule; marginal wing yellowish to stramine-
ous, occasionally tan or brownish, 1 mm. wide, somewhat corky
but scarcely thickened, microscopically foveolate, glabrous (very
rarely with a hair or two), without red spots (very rarely with
one or two) on inner margin; body of achene dark brown, shin-
ing, microscopically latticed, rounded on back, smooth on inner
face or with a few, small, central or scattered, inconspicuous,
callous papillae, and occasionally with some small, appressed,
two-celled, scarcely clavellate or capitate hairs; central sterile
achenes similar to but smaller than fertile ones.

Range. Dry rocky slopes of the inner South Coast Ranges
of California, Santa Clara County to western and central Santa
Barbara County, altitude 150-600 meters; Mount Hamilton
Range, San Carlos Range, southern Gabilan Range, southeastern
Santa Lucia Mountains, western San Rafael Mountains.

ype.) In California legit cl. Douglas.”

222 MADRONO [Vol. 4

Specimens examined. California, Douglas 8, achenes from
type at Herb. Kew. (UC). Santa Clara County: Arroyo Bayo,
Mount Hamilton Range, altitude 2000 feet, April 2, 1936, H. K.
Sharsmith 3490, flowering specimens typical (UC), April 25, 1936,
H. K. Sharsmith 3627, achenes typical (UC), May 15, 1937, H. K.
Sharsmith 3944, 8946 (UC). San Benito County: Hernandez
Valley, May 1, 1933, Roxanna S. Ferris 8394 (UC, SU). Mon-
terey County: Priest Valley, May 13, 1893, Alice Eastwood (UC) ;
Jolon, Bradley road near Bryson road junction, May 2, 19338,
Roxanna S. Ferris 8454 (UC, SU) ; Bradley-Jolon road near Pleyto
road junction, May 3, 1933, D. D. Keck 2098 (P); Jolon, T. S.
Brandegee in 1876 (UC); top of Jolon grade, March 24, 1935,
Alice Eastwood and J. T. Howell 1966 (CA); summit of Mustang
grade, March 26, 19385, Alice Eastwood and J. T. Howell 2085
(CA); Mansfield Ranch, King City, April 1, 1915, Alice Eastwood
4022 (CA). San Luis Obispo County: Paso Robles, May 4, 1926,
Alice Eastwood 13858 (CA). Santa Barbara County: Lompoc,
April 16, 1932, Ralph Hoffmann (SBM); Zaca Mountain, March
25, 19385, D. Avelrod 143, immature (VTM).

The identification of De Candolle’s type of Leptosyne Douglasu
with the South Coast Range species under consideration, and the
necessary transfer to this species of the name C. Douglasi, previ-
ously misapplied to C. californica, has been discussed: under the
latter species. The exact locality in California from which
Douglas collected the type of Leptosyne Douglas is unknown,
but a number of Douglas’ collections indicate that he penetrated
the inner South Coast Ranges, and it is probable that he obtained
it either east of Monterey or north and east of Santa Barbara.
The description of C. Douglasw as given above has been based
upon living plants from the Mount Hamilton Range.

Because of its close alliance with the far more abundant C.
californica, C. Douglasii has remained unnoticed by the very few
California botanists who have collected it. It is important to
realize that the vegetative features which help to distinguish this
species are to be adequately recognized only in the living plants.
In herbarium specimens only the achenal characters stand out as
distinctive (H. K. Sharsmith 3946). It should be noted also that
the younger living plants show the characteristic leaf position,
shape, and glaucous cast far more clearly, the leaves becoming
more terete and flaccid with maturity. To a certain degree the
quite young plants maintain their vegetative characters even
when pressed (Ferris 8394, SU).

Coreopsis Stillmanii var. Jonesit was based on a collection made
May 2, 1882, at Pasadena, California (Marcus E. Jones 3361 pro
parte). Although the wings of some of the achenes do show a
certain amount of corky and rugose thickening suggestive of C.
Stillmanii, varietal relationship to this species is not indicated.

1938 ] SHARSMITH: COREOPSIS 223

On the basis of leaf and floral characters, C. Stillmani var. Jonesiu
is referable either to C. californica or to C. Douglasiu; the achenes
are those of C. Douglasii. Pasadena is far removed from the
range of C. Douglasit as otherwise known. A suspected error
in locality is partially confirmed by the inter-mixture of speci-
mens of C. californica on the two duplicate sheets (P, CA) of the
type collection. Another collection of the same locality and
date (Jones 3373, P) consists of one plant of C. californica. The
suspected error is further confirmed by examination of available
Los Angeles County specimens of the section Euleptosyne; all
are found to belong, on the basis of achenal characters, to C.
californica. In a region as well known botanically as the area
about Pasadena, it is significant that none of the many local col-
lectors has ever reported any species which would correspond
to C. Douglasiti. Mr. Frank W. Peirson, who has made a special
study of the Pasadena flora, states that he has never collected
any Coreopsis with such achenal characters. In consideration of
these facts, it seems necessary to conclude that these Jones Pasa-
dena specimens do not represent a part of the geographic range
of C. Douglasii.

6. CorEeoprsis Stinpmanut (Gray) Blake, Proc. Am. Acad. 49:
342. 1913. Leptosyne Stillmaniti Gray, Bot. Mex. Bound. 92.
1859.

Erect, essentially glabrous annual with several (1-15, mostly
3-5) stems from a tap root: cotyledons 1 cm. long, spatulate,
flattened, only slightly fleshy, spreading: stems yellowish green
to stramineous, leafy on lower half or scapose, sometimes
branched, occasionally with a small, appressed bract on upper
portion, monocephalous, 5-30 (mostly 10-15) cm. tall: leaves
2—10 cm. long, alternate, clustered at base or mostly cauline, only
slightly fleshy, flat, spreading; blade triangular, 0.5—5 cm. long,
often as wide, once or twice pinnate into spatulate, obtuse lobes
1-3 mm. wide and often inconspicuously callous tipped, terminal
spatulate lobe usually broader than the lateral pinnae, blade sim-
ple in depauperate plants; petiole 1-5 cm. long, 1-2 mm. wide,
channelled, sometimes margined with scattered glandular capi-
tate hairs, expanded at base: heads erect, 7-20 mm. high, 1—3.5
mm. wide: involucre barrel shaped, rounded at base; bracts
of outer series 4-8, linear to spatulate, 3-10 cm. long, 1-2.5
(mostly 1.5) mm. wide, obtuse, green or reddish green, flat, not
fleshy, somewhat spreading, angled but not gibbous at base,
prominently bearded at base with long (1 mm.), brownish
glandular hairs, similar but shorter hairs scattered along mar-
gins; bracts of inner series 5—10, ovate, 5-10 mm. long, 2—5 mm.
wide, often lightly angled on midrib, acute with apical tuft of
hyaline hairs, greenish yellow, shining, thin, many nerved, nar-
rowly scarious margined, constricted two-thirds from _ base,

224 MADRONO [Vol. 4

tips spreading in anthesis, connivent in early fruit, spreading in
age: ray florets 5-8, pistillate, fertile; ligule orange yellow, obo-
vate, 5-15 mm. long 3-8 mm. broad, deeply 3-toothed at apex,
spreading horizontally; tube 1-1.5 mm. long, glabrous or with a
few, scattered glandular hairs; style branches linear, obtuse:
disc florets 10-40, perfect, orange yellow, usually all fertile;
tube 1.5 mm. long; annulus glabrous or with a few glandular
hairs; throat expanded, 1.5 mm. long; corolla lobes triangular
ovate, acute, glandular papillate, spreading; anther tips ovate
cordate; style tips triangular acute, glandular papillate; pollen
grains spherical, with short spines: receptacular bracts lanceolate,
5-6 mm. long, 2 mm. broad, obtuse or subacute, hyaline, caducous,
not falling attached to disc achenes: achenes monomorphic, ob-
compressed, obovate, 2.5—5 mm. long, 1.5—2.5 mm. wide; pappus
reduced to a cupule or occasionally with 1-2 short, rigid awns
from rim of cupule; marginal wing stramineous, glabrous, corky
thickened, rugose, microscopically foveolate, erose margined;
body of achene dark brown, smooth but usually not shining,
microscopically foveolate, back rounded and glabrous or with
short, scattered, two-celled, not obviously clavellate hairs which
often arise from callous papillae, inner face with a usually well
developed central row of callous papillae, also with smaller,
scattered papillae bearing short hairs.

Range. Arid foothills on either side of the Sacramento Val-
ley and San Joaquin Valley, altitude 30-900 meters; western
slope of the Sierra Nevada from Butte County to Tulare County,
eastern side of the inner South Coast Ranges in Contra Costa
County, Santa Clara County, and Stanislaus County.

Type. “In the Valley of the Upper Sacramento,” Stillman.

Specimens examined. Butte County: Iron Canyon, May,
1897, Mrs. C. C. Bruce 1987 (P). Placer County: Auburn, April
10, 1865, W. H. Brewer 4520 (UC). Tuolumne County: Coulter-
ville, March 21, 1936, H. L. Mason 11019 (P, UCLA, UC); Chi-
nese Camp, 1937, C. W. Belshaw 2782 (UC). Mariposa County:
Mariposa, April 2, 18938, J. W. Congdon (UC). Madera County:
Raymond, May 9, 1925, Alice Eastwood 12604 (CA); 8 miles west
of Chowchilla, March 21, 1936, R. F. Hoover 821 (UCLA).
Tulare County: April, 1897, J. B. Davy, achenes awned (UC).
Contra Costa County: Antioch, April, 1889, T. S. Brandegee (UC).
Santa Clara County: Seeboy Ridge, May 3, 1935, H. K. Sharsmith
1959 (UC), May 5, 1985, H. K. Sharsmith 8054, achenes awned in
some heads (UC); San Antonio Creek, March 28, 1895, E. J.
Applegate 259 (SU). Stanislaus County: Arroyo del Puerto, Red
Mountains, March 30, 1935, H. K. Sharsmith 1689, April 21, 1935,
H. K. Sharsmith 1818 (UC); Adobe Valley, April 22, 1936, H. K.
Sharsmith 35386 (UC).

Plants of some colonies of C. Stillmanii have leaves entirely

aaa na snd

|
|
\

1938 ] SHARSMITH: COREOPSIS 225

or quite basal, and scapes unbranched and naked (H. K. Shar-
smith 3536), while others show the basal leaves less densely clus-
tered, and the lower part of the scapes with a few cauline leaves.
The extreme of this latter condition is indicated when there is no
basal cluster of leaves, and the branched scapes are strongly
leafy up to half their length or more (Belshaw 2782). These
variations appear, however, not to have any particular geographic
significance. The condition of leafy stems has been used to dif-
ferentiate C. Stillmanit from C. californica which has scapose
stems, but this distinction does not hold.

7. Coreopsis maritima (Nutt.) Hook. Curtis Bot. Mag. t. 6241.
1876. Tuckermannia maritima Nutt. Trans. Am. Phil. Soc. n.
ser. 7: 363. 1841. Leptosyne maritima Gray, Proc. Am. Acad.
(e208. 13868,

Erect, robust, glabrous perennial 38-8 dm. high: stems stout,
hollow, spreading from a thick woody base, much branched
above: leaves alternate, fleshy, 5-25 cm. long, 2—3 times pinnate
into often remote, linear, obtuse, flattened divisions 5-50 mm.
long, 1-4 (mostly 2-3) mm. wide; rachis scarcely wider than
divisions, flattened, extending into a petiole 2-15 cm. long and
2-3 mm. wide at base: heads few (2-3 at ends of branches), 1.5—
2 cm. high, 6-9 cm. wide, erect on essentially naked peduncles
15-50 cm. long: involucre campanulate, rounded and fused at
base into a disc 1 cm. across; bracts of outer series 6-10, oblong
or oblong-ovate, 10-25 mm. long, 4-8 mm. wide, obtuse, green,
flat, spreading to reflexed; bracts of inner series 10-15, ovate,
12-15 mm. long, subacute, thin, yellowish, many nerved, erect or
with tips spreading: ray florets 15—20, pistillate, fertile; ligule
golden yellow, elliptic to narrowly obovate, 25—40 cm. long, sub-
entire to irregularly 3-toothed at apex, spreading horizontally ;
tube 3-4 mm. long, glabrous; style branches linear, obtuse; disc
florets many, perfect, golden yellow, all fertile or central ones
sterile, tube 2.5 mm. long; annulus glabrous or only weakly
bearded with glandular papillae; throat funnelform, 3 mm. long;
corolla lobes ovate, acuminate, thinly glandular papillate within;
anther tips ovate, subcordate at base; style tips triangular acute,
glandular papillate; pollen grains spherical, with short spines;
receptacular bracts linear to oblanceolate, 8-12 mm. long, 1.5—3
mm. wide, subacute, midrib slightly callous thickened, falling
separately from disc achenes: achenes monomorphic, obcom-
pressed, oblong to obovate, 6—7 mm. long, 2-3 mm. wide, plane
or slightly meniscoid, dark brown, smooth or often with micro-
scopic surface papillae, seemingly glabrous but often with micro-
scopic, hyaline hairs, inner face with slight central ridge, back
rounded; marginal wing up to 1 mm. broad, thin, light brown to
dark brown, smooth, microscopically latticed.

Range. Seacoast of southern California in San Diego

226 MADRONO [Vol. 4

County; south along coast to northern Baja California and
adjacent islands, altitude 3-60 meters.

Type. “On shelving rocks, near the sea at St. Diego, in Up-
per California.” Nuttall.

Specimens examined. San Diego County: Del Mar, March
28, 1894, T. S. Brandegee (UC); La Jolla, April 16, 1904, H. P.
Chandler 5123 (UC) ; Point Loma, March 4, 1896, T. S. Brandegee
(UC); Soledad, March 29, 1882, M. E. Jones 3134 (P); San

Diego, March, 1895, T. S. Brandegee (UC) ; Cardiff, May 9, 1924, —

P. A. Munz 7955 (P). Baja California: Salado Cafion, April 27,
1893, T. S. Brandegee (UC); San Martin Island, March—June,
1897, A. W. Anthony 216 (UC).

8. Coreopsis GIGANTEA (Kellogg) Hall, Univ. Calif. Publ. Bot.
38: 142. 1907. Leptosyne gigantea Kellogg, Proc. Calif. Acad.
Sci. 4: 198. 18738.

Erect, robust, glabrous perennial with a thick, fleshy main
trunk 3-30 (mostly about 12) dm. high, up to 1 dm. thick; pri-
mary branches remote, club-like: leaves alternate, clustered at
apices of branches, 3-25 cm. long, 3—4 times pinnate into many
linear to linear filiform, apically truncate divisions 10-50 mm.
long and 0.5—1.5 mm. wide which are lightly grooved above and
rounded beneath; rachis stout, terete, lightly grooved above,
extending into a petiole 30-70 mm. long and 38-5 mm. thick:
heads 1-2 cm. high, 4-8 cm. wide, erect, numerous, cymosely
clustered on somewhat leafy peduncles 6—20 cm. long at the ends
of branches: involucre campanulate, rounded at base; bracts of
outer series 5-12, lanceolate to oblong, 5-20 (mostly 8-10) mm.
long, 2-4 mm. wide, obtuse, flat, green, spreading to reflexed;
bracts of inner series 10-15, oblong ovate, 10-15 mm. long, 4-8
mm. wide, subacute, thin, yellowish, erect, many nerved with
midrib often callous thickened below: ray florets 10—16, pistillate,
fertile; ligule golden yellow, elliptic to narrowly obovate, 20-30
mm. long, 5-8 mm. wide, obtuse or subacute with entire or mi-
nutely and irregularly 3-toothed apex, spreading horizontally;
tube 3-4 mm. long, glabrous; style branches linear, obtuse; disc
florets many, perfect, golden yellow, all fertile, or central ones
usually sterile; tube 2.5 mm. long; annulus glabrous or only
weakly bearded with glandular papillae; throat funnelform, 3
mm. long; corolla lobes triangular acute, thinly glandular papil-
late within; anther tips narrowly ovate cordate; style tips trian-
gular acute, glandular papillate ; pollen grains spherical, with short
spines: receptacular bracts linear, 8-10 mm. long, 1.5—2 mm. wide,
midrib somewhat callous thickened on lower half, falling sepa-
rately from disc achenes: achenes monomorphic, obcompressed,
oblong to obovate, plane or slightly meniscoid, 5-6 mm. long,
2-3 mm. wide, glabrous or with a very few microscopic hairs,
epappose; body dark brown, smooth, shining, microscopically
latticed, inner face with slight central ridge, back rounded:

1938 | SHARSMITH: COREOPSIS 227

marginal wing narrow, thin, smooth, microscopically latticed,
light-brown or dark brown.

Range. Seacoast of southern California from San Luis
Obispo County to Los Angeles County, both on mainland and
adjacent islands, altitude 15-60 meters.

- Type. “Cuyler Harbor, San Miguel Island, about 40 miles
off the coast of Santa Barbara, Cal.,’ W. G. W. Harford.

Specimens examined. Marinianp. San Luis Obispo County:
Oso Flaco Lake, March 17, 1936, H. C. Lee 514 (VTM). Santa
Barbara County: Point Sal, April 11, 1936, Annetta Carter 1095
(UC); five miles south of Surf, April 14, 1929, R. S. Ferris 7572
(P, UC). Ventura County: foothills east of Hueneme, W. G.
Wright in 1894 (UC) ; near Point Mugu, May 14, 1931, C. B. Wolf
2052 (P, UC). Los Angeles County: Malibu Hills, April 26,
1926, M. E. Jones (P); Point Duma, March 7, 1898, J. H. Barber
872 (UC). Istanps. Santa Rosa Island: April, 1901, P. M.
Jones (UC); April 9, 1930, P. d. Munz and E. Crow 11750 (P).
Santa Cruz Island: May 12-15, 1929, L. Ellison (UC) ; vicinity
of Prisoner’s Harbor, April 26, 1930, L. R. Abrams and I. L. Wig-
gins 129 (P, UC). Anacapa Island: May 12-15, 1929, L. Ellison
(UCLA). Catalina Island: Bird Rock, January 22, 1920, C. F.
Millspaugh 4630 (UC); March 27, 1911, 1. J. Condit (UC). San
Nicolas Island: March 18, 1932, J. T. Howell 8220 (P). San
Miguel Island: April 10, 1930, P. A. Mung and J. Voss 11877 (P).
A specimen from Del Monte, Monterey County (May 24, 1923,
Eric Walther, CA) is probably an escape from cultivation.

Discussion

Section Puciopaprus. Although there is a close degree of
relationship among the three species of this section, specific char-
acters are abundant and obvious, and intergradation is quite
lacking, despite the fact that C. Bigelovit and C. calliopsidea oc-
cupy a common geographic territory.

Coreopsis Bigelovit and C. hamiltoni agree in such characters
as scapose stems, basally clustered leaves, and cylindric, truncate
involucre with a variable number of linear outer bracts. Coreop-
sis calliopsidea stands quite apart with stems leafy, lower leaves
obviously clustered, and campanulate involucre with five ovate
outer bracts. When achenes are considered, however, the rela-
tionships are reversed. In C. Bigelovit and C. calliopsidea the
pappus paleae are lanceolate acuminate and the disc achenes
have permanently attached receptacular bracts with calloused
midribs (more obvious in C. calliopsidea). The pappus paleae
of C. hamiltoni are obovate with triangular-acute apices and the
receptacular bracts, which lack the calloused midrib, fall sepa-
rately from the disc achenes. All three species contrast in size
of pappus palea, and in distribution and length of cilia on the
disc achenes (pl. XXXII, figs. 1-17). A striking feature of C.

228 MADRONO [Vol. 4

hamiltonu, not shared by C. Bigelovii or C. calliopsidea, or, indeed,
by any of the Californian species of Coreopsis, is the strongly re-
flexed ligules. The ligules are usually horizontal in the sub-
genus Leptosyne.

When all of the features of similarity and difference between
the three species of the section Pugiopappus are considered, C.
Bigelovii is seen to occupy a position between C. calliopsidea on
one hand and C. hamiltonit on the other. Phylogenetically C.
Bigelovu, or an ancestral type from which it was derived, would
appear to be primitive for this particular unit, C. calliopsidea and
C. hamiltonit having diverged from it in opposite directions.

SecTION EvLeptosyNE. There is much similarity in habit be-
tween the three species of this section, but a closer relationship
is indicated between C. californica and C. Douglasii. In these two
species the stems are always scapose, and the leaves basally
clustered, while in C. Stillmanii there may be great variation in
these characters. Coreopsis Stillmaniti may even equal C. calliop-
sidea of the section Pugiopappus in leafiness and branching of the
stems. Coreopsis californica and C. Douglasi agree, also, in their
fleshy, linear, entire or once pinnate leaves, while C. Stiulmani
has distinctive, flat, spatulate, once or twice pinnate leaves. Less
obvious but no less characteristic leaf differences occur, however,
between C. californica and C. Douglasi, for C. californica has the
leaves erect, linear filiform, terete, and light green, while in C.
Douglasi they are spreading (becoming erect with age), linear,
flattened, and somewhat glaucous. As has already been men-
tioned, the leaf distinctions of C. Douglasi can be observed with
certainty only in young living plants.

The general nature of the barrel shaped involucre is common
to all three species, but in C. californica and C. Douglasi the outer
bracts are fleshy, narrowly lanceolate, and gibbous at the base,
while in C. Stillmanii they are flat, linear to spatulate, and merely
angled at the, base. Sherff relied in part on an extreme varia-
tion in the number of outer involucral bracts for the establish-
ment of C. Stillmanii var. Jonesii. The number of these bracts,
however, is inconstant in all the annual Californian species ex-
cept C. calliopsidea, and this character does not seem to be of
taxonomic significance.

EXPLANATION OF THE FIGURES

Pirate XXXII. Fig. 1. Head of C. Stillmant, X1.5. Fig. 2. Inner face
of achene of C. Stillmanii,x5. Fig. 3. Head of C. californica,x1.5. Fig.
4. Inner face of achene of C. californica,x5. Fig. 5. Head of C. Douglasii, x
1.5. Fig. 6. Inner face of achene of C. Douglasti,x5. Fig. 7. Hair from
achene of CO. californica,X175. Fig. 8. Head of C. Bigelovii,x1.5. Fig. 9.
Inner face of disc achene of C. Bigelovii,x 5. Fig. 10. Inner face of ray achene
of C. Bigelovii,x 5. Fig. 11. Hair from disc achene of C. Bigelovti,x35. Fig.
12. Inner face of ray achene of C. hamiltoniti,x5. Fig. 13. Inner face of
disc achene of C. hamiltonii,X5. Fig. 14. Head of C. hamiltonii,x1.5. Fig.
15. Head of C. calliopsidea,X1.5. Fig. 16. Inner face of ray achene of C.
calliopsidea,X5. Fig.17. Inner face of disc achene of C. calliopsidea, x 5.

1938]

SHARSMITH: COREOPSIS

Puate XXXII. Native ANNUAL CALIFORNIAN SPECIES OF COREOPSIS.

(See explanation of figures on page 228.)

230 MADRONO [Vol. 4

Coreopsis californica and C. Douglasii have the annulus of the
disc corollas obviously bearded, while in C. Stillmanii it is char-
acteristically glabrate. Coreopsis Stillmanii is the only annual
California species of Coreopsis which lacks a distinctly bearded
annulus.

Achenal differences play an important part in species differ-
entiation within the section Euleptosyne as well as in the section
Pugiopappus. One feature is sufiicent to distinguish the achenes
of C. californica, the presence of conspicuous, two-celled, clavel-
late or capitate hairs on body and wing. Even in immature
achenes, with adequate magnification, these minuscule clubs or
stalked knobs stand out clearly along the margin of the wing.
Other characters which aid in distinguishing the achenes of C.
californica are the dull, rough, tan to reddish surface, and the
irregularly thickened, corky wing and central ridge of the mature
and fertile achenes. C. Douglasii is marked by the glabrous
nature of the immature achenes, and the smooth, shining, usually
glabrous, brown body and thin, yellow or stramineous wing of
the mature achenes. Even though the inner face occasionally
bears some callous papillae or two-celled hairs, the hairs are
smaller and scarcely clavellate or capitate, and almost never
(one or two at most) occur on the wing. The sparsely hairy,
smooth, central sterile achenes of C. californica, often with con-
spicuous red spots (“oil glands’), are quite different from the
achenes of C. Douglasii, and are easily distinguished.

The question of specific relationships within this section is
somewhat difficult to visualize. The specific delimitation of C.
Stillmanii is obvious, even though this species transgresses the
separate geographic ranges of both C. californica and C. Douglasii.
Less obvious, however, is the specific differentiation of C. cali-
fornica and C. Douglasii. With the use of herbarium specimens
alone, there is but a single, clear basis for their separation, that
of the achenes, a reliable criterion even when the achenes are
immature. Only in young, living plants are the other differ-
entiating characters adequately seen. A study of simultaneous
garden cultures of C. californica and C. Douglasii, as well as field
observations, gave convincing evidence that additional differences
exist which justify a specific segregation. Another factor of sig-
nificance is that the two species occupy distinct geographic ter-
ritories. The western limit of C. californica is in very close
approximation to the southern limit of C. Douglasii, but, unless
the Pasadena locality of Jones’ specimens should ever be vali-
dated, no specimens are on record which show an overlapping
between the ranges of the two species.

One experiment was performed, which though yielding insuf-
ficient evidence to be significant in itself, may point the way to a
convincing proof of the relationship of C. californica and C. Doug-
lasti. A reciprocal cross was made between these two species

1938 ] BLODGETT AND MEHLQUIST: DELPHINIUM 231

by removing the unopened, perfect disc florets from a young
head of each, and leaving only the pistillate ray florets. The
heads were then cross pollinated. A control was similarly treated
and self pollinated. The heads were protected from insects and
allowed to mature. Fully developed and apparently fertile ray
achenes were obtained from the control, but only imperfect and
partially developed ray achenes were obtained from the recipro-
cal cross.

Coreopsis californica predominates in the area which may be
considered the center of diversity for the subgenus Leptosyne in
California, cismontane and desert southern California. Consid-
ering this fact as well as its wider range and greater abundance,
C. californica may be thought of as a unit ancestral to C. Douglasit.
Whether C. californica or C. Stillmanii should be considered as
primitive in the section Euleptosyne is questionable. On the basis
of distribution, C. californica may rank as the most primitive mem-
ber of the section, but the center of diversity may differ from
the center of origin which often proves of greater significance in
establishing relationships. The occasional occurrence of one or
two smooth, straight awns upon the cupule of C. Stillmanit may
indicate that this species is primitive in the section.

Section TuckerMANNIA. In their perennial habit and mari-
time habitat, the two members of this section are highly distinc-
tive among the Californian species of Coreopsis. They differ
particularly in the arrangement and number of the heads, those
of C. gigantea being numerous and cymosely clustered on short,
leafy peduncles, those of C. maritima being few in number on long,
essentially naked peduncles.

University of California,
Berkeley, July 6, 1937.

COLOR VARIATION IN DELPHINIUM CARDINALE HOOK.
Cuartes O. Biopcetr anp G. L. Mentevistr

Conspicuous variation in flower color within a species in its
natural habitat is of some scientific interest. We wish, therefore,
to record an observation which to the best of our knowledge has
not heretofore been reported.

About four miles southeast of Lompoc, in the northwestern
part of Santa Barbara County, California, there exists an ex-
tensive natural colony of Delphinium cardinale Hook. Through the
kindness of Mr. Ian Sinclair of Bodgers Seed Company we were
privileged to see this colony on July 17, 1937. The field of
approximately eighty acres was on the north slope of a rather
steep, thin-soiled hill, covered with loose calcareous rock from
low cliffs above. The hillside bore a sparse cover of typical
Coast Range chaparral, the shrubs ranging from two to five feet
tall. Scattered among these were many plants of the brilliant

232 MADRONO [Vol. 4

scarlet larkspur, varying in height from three to six feet or more.
The plants were evidently completely at home in this dry sunny
location, sheltered from the stiff cool coast breezes so prevalent
in the valley, for there were many magnificent specimens. Inter-
spersed among the plants bearing the typical cardinal colored
flowers were a conspicuous number with flowers of paler hue,
ranging through several shades of orange-red, orange-yellow, and
buff to a clear lemon-yellow with only a trace of cardinal in the
spurs. The color-variant forms were of course in the minority,
no doubt less than ten percent of the total number of plants, while
the pure yellows, though noticeable, were very few. No other
consistent differences, such as height, habit, or leaf variations,
could be distinguished in the plants.

Similar color variations have been recorded in Delphinium
nudicaule Torr. & Gray, a closely allied Californian species. In
“Delphinium,”’ the book of the American Delphinium Society for
1936, Mr. Carl Purdy (p. 37) mentions such color variations, and
Major N. F. Vanderbilt (p. 65) states that he has “described
various forms and variations in D. nudicaule in a wide range.”
Neither author indicates whether or not the variations were found
in wild plants or in those resulting from garden manipulation.

Division of Genetics, University of California,
Berkeley, October 25, 1937.

GLAUCOCARPUM, A NEW GENUS IN THE CRUCIFERAE
Reep C. Rowwins

The discovery, through recent exploration, of a new genus of
plants in the continental United States is rare enough, it seems to
me, to merit special consideration. It is not surprising that the
more remote parts of the west should yield new species of plants,
but it must be conceded that nearly, if not quite, all the native
genera are known. Thus when a year ago, Dr. E. H. Graham
transmitted to me for identification, specimens of a strange crucif-
erous plant which he had collected in the Uinta Basin of eastern
Utah, the possibility of their belonging to an undescribed genus
seemed remote indeed. An exhaustive study at the time, showed
that these plants possessed a broad relationship with certain
species of “Great Basin’ crucifers which have been variously
considered to belong to Thelypodium, Thelypodiopsis or Sisymbrium.
Graham’s specimens, though only in flower or in some cases pos-
sessing a few immature fruits, revealed distinctive characteristics
which indicated that their disposition in any of the known genera
was unsatisfactory and at best could be only temporary. Mature
fruiting specimens were necessary for a complete analysis of this
anomalous species, hence a trip into the Uinta Basin to make col-
lections and detailed field observations was planned. On June 15,

1938 ] ROLLINS: GLAUCOCARPUM 233

19387, I visited the site of Graham’s collecting and found the
plants growing on a narrow (20 feet) highly calcareous stratum
of shale of the Green River formation. The caespitose plants
growing in clumps of from two to six inches across and deployed
along this single stratum in a highly perplexing fashion, were
traced along the face of a high bluff, known locally as Big Pack
Mountain, for more than three miles.

This close adaptation to a specific stratum combined with the
fact that the plant’s discovery was so very recent, gives some indi-
cation that we are probably dealing with a highly localized
endemic. Further substantiation of this view is evidenced by the
more or less restricted ranges of the species found in association
with it. These include: Yucca Herrimaniae Trel., Linum leptopoda
A. Nels., Mentzelia sp., Gilia polycladon Torr., Cryptantha nana
(Eastw.) Payson, Cryptantha Grahamii Johnston, Cryptantha sp.,
Erigeron argentatus Gray and Hymenopappus lugens Greene. Of
these species, Cryptantha Grahamii is known only from the im-
mediate vicinity, Cryptantha sp. and Mentzelia sp. are apparently
new to science and the collection of Linum leptopoda appears to
be the second for this rare species. The type collection of the
latter was made in southern Nevada. The other species listed
are typically “Great Basin’ and are found most frequently in
eastern Utah and adjacent Colorado. The deployed distribution
and paucity of individuals of all species in the limited area under
consideration, gives evidence of the extremely dry conditions
under which they survive. The area is in the Upper Sonoran
Life-Zone and would ordinarily be placed in the mixed desert
shrub type, although the actual site more nearly approaches a
“bad land” type of habitat.

Glaucocarpum gen. nov. Perenne suffruticosum glabrum et
glaucum; caulibus gracilibus simplicibus; foliis alternis integris
vel sparse dentatis; inflorescentiis racemosis; sepalis oblongis non
saccatis; petalis flavis spathulatis integris; pedicellis erectis
rigidis; siliquis sessilibus vel stipitatis glabris et glaucis; stylis
robustis ; stigmate integro ; loculis 4—8-ovulatis ; seminibus oblongis
exalatis uniseriatis ; cotyledonibus incumbentibus.

Glaucocarpum suffrutescens (Rollins) comb. nov. Species
typica. Thelypodium suffrutescens Rollins ex Graham in Ann.
Carneg. Mus. 26: 224. 1937. The known collections are from
the Uinta Basin of eastern Utah and include the following: west
of Willow Creek, Thorne’s Ranch, eastern slope of Big Pack
Mountain, Uintah County, May 23, 1935, Graham 8950 (type in
Gray Herb., isotypes in Carnegie Mus. Herb.); June 15, 1987,
Rollins 1700 (Gray Herb.).

The study of a large series of fruiting plants both in the field
and as specimens, make advisable certain minor changes and ad-
ditions to the original description of G. suffrutescens. Plants
caespitose, 1-2.5 dm. high (average 2 dm.) root strong, deep and

234 MADRONO [Vol. 4

with a pithy texture; radical leaves absent; leaves entire or
sparsely and remotely dentate, petiolate or sessile by a narrow
base, elliptical to broadly oblanceolate; paired stamens united
at base or appearing as a single stamen as a result of the uniting
of filaments and anthers by their entire length; siliques 1-2 cm.
long, 2-8 mm. broad, slightly flattened parallel to the septum,
sessile or with a short, stout gynophore less than 1 mm. long;
valves strongly nerved from base to apex; style stout, 1-2 mm.
long; stigma circular, entire and unexpanded; ovules 4-8 in each
cell; seeds uniseriate, oblong, plump, 1.5—2 mm. long, 1-1.5 mm.
broad, mucilaginous when wetted; funiculus free, stout, less than
1 mm. long; septum with a median band of cells slightly
elongated parallel to replum; cotyledons incumbent.

Glaucocarpum is most closely related to certain species of
Thelypodium, which in the broader sense includes species placed in
Thelypodiopsis by Rydberg (Bull. Torr. Bot. Club 34: 432. 1907)
and by O. E. Schulz (Engler, Pflanzenfam. 17b?: 582. 1936)
and in Sisymbrium by Payson (Univ. Wyo. Publ. Sci. 1*: 11-18.
1922). However, the natural relationship is not sufficiently close
to allow the inclusion of the new plant in any of these genera. In
order to avoid confusion in making a comparison, the broadly
defined ““Thelypodium” sensu Robinson (Syn. Fl. N. Am. 1*: 173.
1895) and Jepson (FI. Calif. 2: 35. 1986) is adopted for the
present discussion, but it is not necessarily accepted as the most
natural treatment. At the outset, Glaucocarpum and Thelypodium
are dissimilar in field aspect. Except for being more caespitose
and of lower stature, the new genus has a closer resemblance to
Stanleya in habit and general appearance than to any species of
Thelypodium. Distinctive features of Glaucocarpum are: the
deep, strongly perennial, woody root; branching multicipital
caudex; slender, wiry, simple and leafy stems; absence of radical
leaves; numerous petiolate or narrowly sessile, entire cauline
leaves; narrow, strict and elongated but lax inflorescence; green-
ish-yellow petals and young sepals; united long stamens; slightly
developed nectar glands; strongly nerved, broad, rather short
siliques and stout evident styles. Opposing characteristics typical
of the genus Thelypodium include: a biennial or short-lived peren-
nial habit; simple caudex; stout branching or more rarely simple
stems; well-differentiated radical leaves; cauline leaves which
are sessile and auriculate or if petiolate, then divided; broad and
usually very congested inflorescence; purple to white or rarely
yellowish flowers; long narrow often apiculate anthers with free
filaments; well-developed nectar glands; long narrow and nerve-
less or nerved siliques with a short or obsolete style. In the
aggregate, these characters effectively demonstrate the generic
distinctiveness of Glaucocarpum.

Gray Herbarium, Harvard University,
February 18, 1938.

1938 | ROLLINS: GLAUCOCARPUM 235

Pirate XXXIII. Giaucocarpum suFFRUTESCENS Rouuins. Fig. 1. Fruiting
plant X% (Rollins 1700). Fig. 2. Flowering raceme from the type x2
(Graham 8950). Fig. 3. Siliques x3 (Rollins 1700).

236 MADRONO [Vol. 4

REVIEWS

Tiibatulabal Ethnography. By Erminize W. Vorcerin. Anthro-
pological Records. Volume II, Number 1. Pp. 1-84 with 6
plates and 16 text figures. University of California Press.
March 9, 1938. $1.00.

Mrs. Voegelin has made an ethnographic study of the Kern
River Indians of California during which extensive observations
were made of the use by these primitive peoples of the native
plants at their disposal. Specimens were made of the plants
used and the collection deposited at the University of California
Herbarium. Of 220 plant species collected by the author, 130
are reported to be used as food or medicine or are applied in
some way in the primitive handcraft of the Indians. The native
names for most of the plants are recorded. Considerable space
in the published report is devoted to an account of the tending
and the preparation for use of native tobacco. Both Nicotiana
Bigelovit Wats. and N. attenuata Torr. were used. A summary of
the reported use of tobacco by several Indian tribes is made.
Another section deals with the collection, preparation and storage
of plant foods. Acorns and pifion nuts are among the most im-
portant of these. In addition to the ethnobotany that is defi-
nitely organized, there is much casual reference to plants and
plant uses that would interest the botanist searching for such
information. The literary style is telegraphic. The printing is
in photolithography. It is gratifying to see that such informa-
tion is being gathered and preserved before it becomes lost.—
Hersert L. Mason.

Species Lupinorum. By C. P. Smitu. Privately published by
the author, Route 1, Box 24, Saratoga, California. Signature 1,
pp. 1-16, April, 1988. Signature 2, pp. 17-32, May, 1938.

Botanists will welcome the first two numbers of the “Species
Lupinorum”’ by Charles Piper Smith. The projected work is the
culmination of the author’s years of experience with the difficult
genus Lupinus and will include the lupines of the world. Signa-
ture one of April, 1938, comprising a part of paper one, includes
besides the introduction, a “Catalog of the published names pro-
posed for species,’ a “Catalog of names of authors describing
one or more species or varieties,’ and the beginning of a ““Chrono-
logical catalog of species and varieties” closing with the year
1910. The second signature, May, 1938, continues the chrono-
logical catalog to the year 1987, thus closing paper one. Paper
two of May, 1938, also included in the second signature, is “A
preliminary catalog of the lupines of Baja California and Sonora.”’
The species and varieties and their synonyms, the type collec-
tions, bibliographic references, and the citations of a few speci-
mens calculated to indicate the range are listed. No descrip-

ets

1938 | NOTES AND NEWS 237

tions occur except for new species and varieties. Where necessary,
critical notes are included to clarify taxonomic problems. Lu-
pinus Mearnsii C. P. Smith, L. albifrons var. Brandegeei C. P. Smith,
L. latifolius var. Wigginsii C. P. Smith are described as new. The
paper closes with a key to the species included. It is to be hoped
that nothing will stand in the way of the completion of this very
worth while series of papers. The printing is in photolithog-
raphy. Copies may be had by making a contribution to the pub-
lishing fund. A minimum contribution of one dollar entitles the
donor to five signatures.—HeErBert L. Mason.

Plants of the Lava Beds National Monument, California. By
Ermer I. Appriecate. The American Midland Naturalist.
Volume XIX, Number 2. Pp. 334-368. March, 1938.

So little has been written of the western extension of the
Great Basin flora into the Pacific Coast states, that this capable
treatment of one such area is extremely valuable. The Lava
Beds National Monument comprises about seventy-two square
miles of northeastern Siskiyou County and adjacent Modoc
County. The numerous cinder-cones, lava flows and deposits of
pumice, which support a rather sparse vegetation, clearly mark
it as a region of recent volcanic activity. The author lists ten
species of trees, thirty three of shrubs, one hundred and fifty of
herbs, and two of ferns, many of which are typical of the Great
Basin, although some are evidently derived from the Sierra-Cas-
cade and Klamath (Siskiyou) areas. <A well written, popular
account of the “‘life zone’’ concept, and terse keys to the families,
genera and species should make the paper helpful to interested
amateurs as well as to professional botanists.—Lincotn Con-
STANCE,

NOTES AND NEWS

BaTTAREA PHALLOIDES (Dicks.) Pers. In Santa Barzpara. This
stalked puffball, although widely distributed, has not heretofore
been reported from Santa Barbara. It was observed in rocky
soil near the seashore, January, 1938. Dimensions noted below
exceed those usually given in descriptions of this species: stem
hollow, woody, shining straw color, 25 cm. long, 2.5 to 3.5 cm.
in diameter, tapering toward the cap; stem scales conspicuous,
pendulous, 5 to 10 mm. wide, 10 to 25 mm. long; volva subter-
ranean, 10 cm. in diameter with loose ragged edge; cap bell-
shaped, 7 cm. in diameter, 5 cm. in depth, with smooth, white
skin-like peridium which at maturity splits horizontally along
the edge and is gradually pushed off; exposed mass of red-brown
spores gradually disappearing, the stem standing for several
months.—RutH Hartweitt, Museum of Natural History, Santa
Barbara, California.

238 MADRONO [Vol. 4

Dr. W. H. Long, Senior Pathologist, retired, United States
Forest Pathology Field Office, Albuquerque, New Mexico, was a
visitor, during May and June, to the herbaria at Stanford Uni-
versity and the University of California, Berkeley. Dr. Long was
especially interested in studying the collections of gasteromycetes
and rusts, particularly Ravenelia.

The members of the Second Expedition of the University of
California Botanical Garden to South America will sail from San
Francisco for Peru on July 20,1988. The staff, under the direc-
tion of Dr. T. H. Goodspeed, will include Dr. H. E. Storck, Carle-
ton College, C. R. Worth, Rutgers University, A. A. Beetle,
Rocky Mountain Herbarium, W. J. Eyerdam, Seattle, Washing-
ton, J. L. Morrison and O. B. Horton, University of California,
Berkeley. The expedition was organized for the purpose of col-
lecting plants suitable for garden culture as well as for obtaining
herbarium specimens. On reaching Peru, the members of the
expedition will divide into several parties, one covering the higher
portions of Peru and Bolivia, another, the Peruvian and Chilean
coasts, and a third, southern Patagonia and Tierra del Fuego.

Miss Sarah C. Dyal, on leave from the Herbarium, Depart-
ment of Botany, Cornell University, has been making a field study
of the Valerianaceae, mainly Plectritis, during the past year.
Most of April and May, 1988, were spent in central California,
where in addition to taking numerous field trips, she visited the
herbaria at Stanford University, California Academy of Sciences
and University of California. After leaving Berkeley, Miss Dyal
went north along the coast, stopping in Eureka to consult the
private herbarium of Joseph P. Tracy.

Of especial interest to western American botanists is the re-
cent appearance of the “Flora of the Aleutian Islands and west-
ernmost Alaska peninsula with notes on the flora of the Com-
mander Islands,” by Eric Hultén (pp. 1-397. Stockholm, 1937),
Curator of the Herbarium at the Botanical Museum of the Uni-
versity of Lund, Sweden. The introduction deals with the his-
tory of botanical collecting as well as the geological history and
phytogeography of the area. The taxonomic portion of the
volume consists of 298 pages of discussion, literature citations,
and ranges. Descriptions are given only for new species and
varieties. Geographical ranges of the 477 species treated in the
flora are summarized in a series of maps at the end of the volume.

Mrs. Ynez Mexia has recently returned from Mexico where
she has been collecting in the States of Guerrero and Oaxaca
since last October. In the latter state she worked in the vicinity
of Mitla and Yaveo, a region little known to botanists.

1938 ] PROCEEDINGS 239

Dr. Carl Epling, Department of Botany, University of Cali-
fornia at Los Angeles, made a trip up the Redwood Highway of
California during June in order to study the Labiatae in that
region.

The following recent publications are of interest to botanists
of western North America.

“Notes on the Flora of the Charleston Mountains, Clark
County, Nevada,” by Ira W. Clokey (Bull. So. Calif. Acad. Sci.
87: 1-11. 1938). Mr. Clokey, who has been making extensive
collections in the Charleston Mountains for several years, de-
scribes six new entities in this paper: Calochortus rhodothecus,
Aquilegia scopulorum subsp. perplezans, Potentilla cryptocaulis, Di-
taxis diversiflora, Angelica scabrida Clokey and Mathias, Senecio
Andersonii.

“Studies in Penstemon VI. The section Aurator” by David
D. Keck (Bull. Torrey Bot. Club 65: 233-255. 1988). In this
paper, the section Aurator is erected and the following subspecies
and varieties are proposed as new: Penstemon Jamesii subsp. brevi-
culus, P. eriantherus var. grandis Pennell and Keck, P. eriantherus
var. redactus Pennell and Keck, P. Whitedw subsp. tristis Pennell
and Keck.

“Plants of Rock Creek Lake Basin, Inyo County, California, »
a check list” by Frank W. Peirson, privately published by the
author, 1988. Mr. Peirson has collected and studied the plants
of this region intensively for a number of years. The remark-
ably complete annotated list contains over 320 species and varie-
ties found above 10,500 feet elevation in this area on the east side
of the Sierran crest.

PROCEEDINGS OF THE CALIFORNIA
BOTANICAL SOCIETY

The California Botanical Society celebrated its Silver Jubilee
on the evening of April 23, 1938, on the occasion of the annual
dinner. This was held in the Berkeley Women’s City Club and
was attended by about 130 members and friends of the Society.
Special effort was made to reach as many as possible of the re-
maining 205 charter members of the Society, for the Silver Jubi-
lee was planned to honor them and particularly the founder of
the Society, Professor W. L. Jepson.

_ The entire second floor of the club house was at the disposal
of the Society for the evening and many old acquaintances were
renewed as friends came together who had become active in other
than botanical lines in recent years. Amateur and professional
botanists were there in equal numbers.

Professor W. W. Robbins, of the University of California
Agricultural Experiment Station at Davis, presided ably as toast-
master following the dinner. He was introduced by the Presi-

240 MADRONO | . [Vol.4

dent of the Society, Professor H. E. McMinn. Dr. Robbins called
for short speeches from several persons including Mr. James B.
Smith and Mrs. Adeline Frederick, charter members, who com-
bined early reminiscences with tributes to the founder; Profes-
sor W. W. Mackie, who discussed some of the early field trips of
the Society and the interesting personalities who attended and
inspired them; and Professor George J. Peirce, past president of
the Society, who read a series of communications from charter
members who were unable to attend the banquet, yet who wished
to recall the inspirational leadership, good comradeship and
steady progress that had marked the organization since its found-
ing. Dr. Peirce then announced the election of Dr. Jepson to
life membership in the Society and called upon him as the last
speaker. |

Dr. Jepson, in addressing the large group of friends and early
associates, emphasized the ideals upon which the California Bo-
tanical Society was founded and which guided its early leaders.
He concluded with an exhortation to the coming ranks to adhere
to the ideals of the Society in making new advances. After touch-
ing upon the early meetings and the labors of many to insure the
success of the organization, the speaker related the historical
background of the founding of the Society. One of the functions
of the California Botanical Society as visualized by its founder
was realized during the first year of the organization, when the
members of the Second International Phytogeographic Excursion
were entertained at dinner at the Hotel Oakland, Sept. 12, 1913.
This occasion brought together perhaps the most notable assem-
blage of world-renowned botanists in the history of the state.

The history of Madrofio was related by the speaker together
with the reasons for its name and format. The journal has al-
ways been edited as an inviting magazine to the reader, with
diversity of content, accuracy and dignity.

Dr. Jepson then suggested means by which the needs of West
Coast botanists may be more fully met by the Society. These in-
cluded the organization of long distance field trips to outstanding
botanical localities, such as the Mohave Desert and the ‘‘White
Plains” or pine barrens of the Mendocino coast. He pointed out
the desirability of preparedness to aid in preserving and conserv-
ing our botanical heritage when such occasions arise, as when, for
example, the largest known individuals of our tree species are
threatened with destruction.

After the address the gathering lingered for social inter-
course and the opportunity to greet the founder. A suitably in-
scribed herbarium sheet was passed around for “‘annotating” and
the entire party enrolled thereon, following which the document
was presented to Professor Jepson.—D. D. Keck.

ide

Complete Your Files!

MADRONO

A West American Journal of Botany

“Madrono” is a quarterly journal de-
voted to important and stimulating ar-
ticles dealing with plant morphology,
physiology, taxonomy, and botanical his-
tory, as interpreted and recorded by west-
ern American botanists. These volumes
should be a part of every botanist’s li-
brary and should be made accessible to
students of all universities and colleges.

Volume I, 1916-1929. . . $5.00
Volume II, 1930-1934 .. 5.00
Volume III, 1935-1936 . 5.00
Simele numbers .).\./.\.).". 0.75

There is only a limited supply of Vol-
umes I and II.

Address all communications
and orders to:

Dr. David D. Keck

Carnegie Institution of Washington
Stanford University, California

NUMBER 8

oot

.
maki
i

A WEST AMERICAN JOURNAL OF
BOTANY

ve

Contents

Tue FLoweriInc oF WOLFFIELLA LINGULATA (Hecetm.) Heceim., Herbert

GPP TEES OREO Neve os A, CUAL NR ELBE Wn Sel 2M 0 GDR A UU 241
THe WESTERN AMERICAN SPECIES OF Parconsa, G. Ledyard Stebbins, Jr. .. 252
Hanoine GARDENS OF THE Canary Istanp Date Pato, Ira L. Wiggins .... 260
ADDITIONS TO OUR KNOWLEDGE OF THE F'torna oF Mount Baker, WASHINGTON,

eG EC TESCHVOR aE ener NN UI Rta nil taste Ue NOR US ROD CNL UA A Hr 263
Notes on Stacuys ricipa Nort., Carl Epling ..............0.. 200000000. 270
Eremocarpus BENTHAM: Preoccupiep ? Louis C. Wheeler ............... 272
SES NEES re EP. PB LACED olf Se eee Uo ete 273
VisE Feniciratis: THe BEGINNING YEARS OF THE CALIFORNIA BOTANICAL

IGEN VN Bes) P28, OPSOM, oii ie hls Se ae ale a I Oh SE 276
Notes oN THE GeNvs Rises In Catirorntia, Clarence R. Quick ............ 286
A Hysrem Erxioconum, Herbert L. Mason .............. 0.0000 290
USSORSES Ra Moet ons a A I) AE Ne ae i Ue HTS oe RE po 291

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October, 1938

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

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Frontispiece to Vol. 4, Maprono.

1938] MASON: FLOWERING OF WOLFFIELLA LINGULATA 241

THE FLOWERING OF WOLFFIELLA LINGULATA
(HEGELM.) HEGELM.

Hersert L. Mason

The occurrence of flowering in the Lemnaceae has been a sub-
ject of great interest during the past ten years. Most of these
reduced aquatic plants either flower rarely, or more probably
because of their minuteness and obscurity, the flowers are easily
overlooked. Flowers in certain species of Lemna and Wolffa, in
some localities at least, are not infrequently observed, while those
of Spirodella are rarely encountered. Until 1935 flowers of Wolff-
ella were unknown although members of this genus are wide-
spread throughout the warmer parts of the earth and most of the
species have been known for approximately seventy years. Be-
cause of the failure to find any flowering specimens during this
long period, Wolffiella has been regarded by some as the most
specialized member of the family, having lost completely its
capacity to flower. It was supposed that sexual reproduction
did not exist and the plants were regarded as being dependent
upon vegetative budding as a means of reproduction. Hicks
(6) in dealing with this subject states (p. 116-117), ‘‘The ability
to produce flowers apparently has been so completely lost that
probably they are never produced by plants in nature. In Wolff-
ella floridana, at least, it is doubtful as to whether the flowering
potentiality could be made to find expression as the result of
favorable physiological condition.” That Wolffella still retains
the capacity to flower is borne out by the finding of flowering
material of W. oblonga (Phil.) Hegelm. in Argentina and of W.
lingulata (Hegelm.) Hegelm. in California.

The first species of Wolffiella to be reported flowering was W.

oblonga, a species ranging from South America to North America.

Flowers of this species were observed by Giardelli (1) in a la-
goon near the town of Dolores, Province of Buenos Aires, Argen-
tina. Careful dissections were made and a very complete diag-
nosis of the floral characters was drawn up. | Illustrations were
presented to supplement the descriptions.

In June, 1937, flowering specimens of W. lingulata were found
in a slough of the marshes of Roberts Island in the delta of the
San Joaquin River near Holt, California. On January 11, 1988,
a collection of sterile plants was made in Trapper’s slough on
Roberts Island. These were brought into the laboratory and
grown in an aquarium. In three weeks flowers began to appear,
and blooming continued over a period of six weeks. In June,
1938, flowering plants were encountered in great abundance at
the Roberts Island locality. From these plants seeds developed
which germinated freely. On August 26, 1938, many plants were
still flowering, although not in such abundance as in June.

Maprono, vol. 4, pp. 241-308. October 31, 1938.

WOV - 2 1938

242 : MADRONO [Vol. 4

W olffiella lingulata occurs in quiet water of sloughs in clearings
among tules (mostly Scirpus acutus). The sloughs are largely the
result of excavation for materials to build the dikes which with-
hold the waters of the San Joaquin river from the rich agri-
cultural delta lands. They have no outlet and obtain their water
through seepage from the diked lands as well as from the river.
The underlying soil is an immense peat deposit. Associated with
the Wolffiella were Riccia fluitans, Typha latifolia, T. angustifolia,
Scirpus acutus, Spirodella polyrhiza, Lemna minor, L. cyclostasia,
Ceratophyllum demersum, Myriophyllum hippurioides, and Jussiaea

californica. Both species of Lemna were flowering throughout the

summer months.

Wolffiella lingulata was described by Hegelmaier (3) from
material collected by Hahn near Mexico City in 1868. A very
complete and revised description is given by Thompson (9)
which, in certain details, is at variance with the observations
recorded here. The vegetative organography has been very ably
discussed by Goebel (2). In general this species possesses a
very simple rootless thalloid plant body (frond) with a triangular
vegetative reproductive pouch at the basal end (pl. XXXVI, fig.
2). The plants are free floating in water at or below the surface
but usually do not break through the surface tension layer until
the time of flowering. Field observations throughout the entire
period of growth and reproduction demonstrate considerable
seasonal variation in the size and shape of the fronds as well as
in the behavior of the plants. There appear to be two well
marked phases of the plant depending upon whether it is in the
vegetative or the flowering condition. As in other members of
the Lemnaceae the flowering phase in Wolffiella is smaller than
the vegetative phase (pl. XXXVI, fig. 3). In Wolffiella, more-
over, it is accompanied by an asymmetrical widening of the basal
portion of the frond to accommodate the floral cavity in which
the inflorescence is produced. Apparently a reduction in the size
of the daughter fronds produced takes place quite generally in
the colony just prior to the time of flowering. That there is a
connection between flowering and size of the fronds is indicated
by the fact that in material growing in the laboratory as well as
that in the field, the average size of fronds in colonies producing
flowers is much smaller than in those reproducing only vege-
tatively.

The frond varies from broadly oblong to linear and may range
from apparently symmetrical to strongly falciform. There is
great variation in proportion of length to width. The length
ranges from 1.5 times the width to as much as 7 times the width.
Often those that are short have the greatest actual width. The
maximum width occurs on vegetative plants. They range from
3.5 millimeters to as much as 5 millimeters wide. Fronds with
lengths of 8 or 9 millimeters are not uncommon. Flowering
fronds however may be from 8 to 5 millimeters long and as nar-

1938] MASON: FLOWERING OF WOLFFIELLA LINGULATA 243

row as 1 to 1.5 millimeters. Much of this variation in size and
shape is seasonal, and descriptions based on a collection taken at
any particular season may be very misleading, particularly when
applied to a plant from the same locality collected at another
season of the year. Except in the flowering condition all of the
plants are concave dorsally by virtue of the turning up of the
lateral margins. In addition the ends of the fronds are projected
downward and only a small portion of the surface area of the
plant ever breaks through the surface layer of the water. This
curvature of the frond causes it to appear as though it were a
“segment of a band.” Some of the fronds may also be twisted
so as to appear somewhat spiral. The vegetative phase seems
to be at its highest development in California during the winter
months. At this time the fronds are of maximum size and show
the greatest variation.

The most common type of reproduction encountered in Wolff-
ella is vegetative and is accomplished by budding of the meriste-
matic tissue at the inner angle of the reproductive pouch. The
young bud is at first strictly symmetrical and is attached to the
parent by means of a stipe (pl. XXXV, figs. 7, 8). Goebel re-
gards this as a very degenerate vascular strand. The length to
which the stipe will develop shows great variation when corre-
lated with season. Early in the ontogeny of the bud, tissue dif-
ferentiation takes place and there is formed at one side of its
stipe axis a triangular reproductive pouch with its opening facing
the parent frond (pl. XXXV, fig. 7). This pouch causes an
asymmetrical development of the new frond with respect to its
axis as indicated by the position and direction of the stipe. The
pouch lies either to the right or the left of the axis. As the
frond continues to grow it assumes an apparently symmetrical
form, but morphologically the axis is diagonal and follows along
one margin of the reproductive pouch. In plants with short
stipes the growth of the daughter frond soon causes it to break
away from the parent. A definite abscission layer made up of
several layers of transverse cells forms across the stipe (pl.
XXXV, fig. 8). During the winter months, however, the stipes
are very long and often the daughter fronds remain attached to
the parent, forming what have been termed family colonies (pl.
XXXV, figs. 4, 5). This is contrary to the observations of
Thompson who states that in W. lingulata family colonies never
occur. In material observed during January as many as ten
pairs of fronds were seen attached in a family colony each with
its stipe sufficiently elongated to accommodate it to colonial exis-
tence. Thompson’s failure to observe colonies may well be due
to the fact that he was dealing with a single seasonal variant with
short broad fronds of large proportion. Such size and shape is
a definite obstacle to the formation of family colonies. Of this
short, broad type of frond not more than five were noted in one
family and these families were rare.

244 MADRONO [Vol. 4

The position of the stipe scar on the frond has been used as
a diagnostic character for this species of Wolffiella. Careful ob-
servation of large numbers of plants demonstrates that the stipe
scar may be either on the lower margin of the reproductive pouch
to the right or to the left of the median axis of the frond or it
may be in the exact right or left corner of the frond (pl. XXXV,
figs. 9, 10). Both dextral and sinistral fronds are observable in
the photograph.

The tissue of the frond is very simple in its structure. It is
only two cells thick at the distal end, whereas at the basal end in
the region of large intercellular spaces it is several cell layers
thick. Such intercellular spaces are characteristic of members
of the Lemnaceae. In Wolffiella lingulata they occupy only the
basal third or half of the frond (pl. XXXVI, fig. 2). The epi-
dermal layers are beset with scattered dense cells which have
been termed pigment cells. Upon drying these become red
brown. The few stomata (pl. XXXV, fig. 6) that have been ob-
served were on flowering specimens and only on that portion of
the frond that is emersed. Goebel reports his inability to see
stomata. Hegelmaier noted one or two near the margin of a
frond. Although search was made no stomata were seen on
strictly vegetative plants. This coincides with the findings re-
ported by Giardelli for W. oblonga. Whether or not the develop-
ment of stomata precedes the emergence of the frond from water
has not been observed. In any event just prior to flowering the
fronds break through the tension layer of the water surface and
soon the development of the inflorescence becomes evident.

The inflorescence occurs in a floral cavity and makes its ap-
pearance first as a double colorless spot composed of a dense
mass of very small cells on the side of the reproductive pouch
that bears the stipe of the frond (pl. XXXVI, fig. 3). Soon it
loses its double aspect and appears as a single oblong mass in a
floral cavity lying in a plane essentially parallel to the side of the
vegetative pouch. Next a slit-like opening occurs in the surface
of the frond above the cavity; this is followed by the emergence
of the single flask-shaped pistillate flower. Flowering is thus
protoginous. When the stigma becomes receptive a small glob-
ule of liquid is exuded (pl. XXXVI, fig. 4) which assumes a
spheroid form and completely covers the concave stigmatic sur-
face; presumably this is a trap to catch insect or wind borne
pollen. With the disappearance of the globule of liquid from
the stigma, the staminate flower, composed of a single stamen,
begins to emerge. The stamen, lying back of the pistillate flower
away from the base of the frond, is made up of a subspheroid
two-lobed anther on a stout filament. During its development
the stamen usually tears the aperture of the cavity. Soon after
emergence it dehisces its dry white powdery pollen and appears
as a conspicuous glistening white spot on the surface of the frond

Prare XXXIV. Worrrrerta trncurata (Hegelm.) Hegelm. Group of
flowering plants, <5. The shaded portions of the fronds are immersed, the
lighter portions bearing the flowers are emersed. (1) Sinistral frond bearing
a dehiscing stamen; (2) dextral frond with the pistillate flower in anthesis
(dextral and sinistral fronds being interpreted on the basis of the position of
the stipe scar rather than the position of the flower).

1938] MASON: FLOWERING OF WOLFFIELLA LINGULATA 245

(pl. XXXIV). It then withers and disappears. There is no evi-
dence of a spathe surrounding the inflorescence.

The young fruit develops asymmetrically into an indehiscent
utricle with a single ellipsoid seed and a persistent style (pl.
XXXVI, fig. 9). In all plants observed the ripening of the fruit
is accompanied by the death of the parent frond, which remains
floating with the fruit in place in the ruptured floral cavity (pl.
XXXVI, fig. 12). Soon the seed begins to germinate. The
ovary wall collapses and disintegrates. The minute brown coni-
cal operculum of the testa begins to push out, attached to the
hypocotyl (pl. XXXVI, figs. 18-18). As the hypocotyl further
enlarges, the operculum becomes increasingly lateral in its posi-
tion and an irregular slit appears separating the hypocotyl from
the cotyledonary sheath. From this slit, the young vegetative
frond makes its appearance, the distal end of the frond emerging
first. The cotyledon remains within the testa. When fully
formed the young plant breaks away from the hypocotyl and
other remaining embryo tissues and is a free floating plant with
the next vegetative generation already formed in its pouch (pl.
XXXVI, fig. 18). At the lower margin of the vegetative pouch
the minute stipe by which the young plant was attached to the
hypocotyl] is clearly visible. A formal description of the inflores-
cence and flowers of this species follows.

WoLFFIELLA LINGULATA (Hegelm.) Hegelm. in Engler, Bot.
Jahrb. 21: 803. 1895. Wolffia lingulata Hegelm. Monogr. Lem-
nac. 182. 1868. Wolffella oblonga of California authors, non
Wolfiella oblonga (Phil.) Hegelm. 1857.

Fronds monoecious, protoginous; inflorescence without spathe,
borne in floral cavity in dorsal side at basal end of frond, dex-
trally or sinistrally to vegetative pouch; staminate flower pos-
terior to pistillate flower, stamen one, filament stout, .74—.76 mm.
long, anther subspheroid, two lobed, white; pollen white, sub-
spheroid 20-23 microns in diameter, minutely mucronulate; pis-
tillate flower, solitary, pistil flask shaped, .47—.48 mm. high, ovary
.13—.20 mm. wide, one-celled, style short, thick, abruptly expand-
ing to a concave circular stigma; ovule solitary, suberect, becom-
ing tilted; fruit a utricle, bladdery, asymmetrical, indehiscent,
style persistent; seed ellipsoid ovoid, .41—.44 mm. long, .29 mm.
wide, glistening white, operculum of testa lying in a cavity at end
of seed (pl. XXXVI, fig. 11).

Specimens from the Missouri Botanical Garden (M) and from
the Herbarium of the University of California (UC) were
examined.

Mexico. Near Mexico City, L. Hahn (type collection, M).
Catirornia. Kern County: near Bakersfield, Oct. 7, 1895, C. H.
Thompson (M). San Joaquin County: one-half mile southwest
of Holt, H. L. Mason 11,548 (UC); Trappers Slough, Roberts
Island, H. L. Mason 11,850, 12,072 (UC). San Bernardino

246 MADRONO [Vol. 4

County: near San Bernardino, Parish 4581 (UC); San Bernardino
Valley, Parish 4586 (UC). Orange County: San Juan Capistrano,
Abrams 4200 (UC). San Luis Obispo County: Oceano, Nov. 14,
1908, I. J. Condit (UC). Monterey County: 2 miles south of
Pajaro, Sept. 27, 1903, C. H. Thompson (M). Santa Clara
County: Alviso, Sept. 9, 1903, C. H. Thompson (M).

Comparing the above description with that given by Giardelli

for Wolffiella oblonga the two species are strikingly alike in flower ©

characters. Only two points of difference are outstanding; a
third may or may not be significant. For W. oblonga there is re-
ported pollen 11 to 15 microns in diameter whereas in W. lingu-
lata the pollen is 20 to 23 microns in diameter. In W. oblonga
the pollen is covered with minute wrinkles whereas in W. lingulata
it is turgid and minutely mucronulate. In general the size of all
the flower parts in W. lingulata exceeds that in W. oblonga. This
is consistent with the larger plant body. The following is a de-
tailed comparison of W. oblonga with W. lingulata as to points in
which measurements were recorded by Giardelli. Except as

noted above and in the following table, the flowers agree in other

characters. The size of the frond was computed from the mag-
nification cited on Giardelli’s illustration.

W. oblonga W. lingulata
Bollene on soci ee es 11-15 mu 20-23 mu
aS tHN ic. ove techn tenet reat 4 mm. high A7—A8 mom. high
.2 mm. wide .13—.20 mm. wide
UI. cee Gee one eee A5— .55 mm. long .58-.77 mm. long
37— 46 mm. wide 45-.50 mm. wide
Seed vile eae enue 35-— 40 mm. high 41-44 mm. high
.25— .29 mm. wide .29 mm. wide
Frond). S ies eee ie nee 2.8 -3.5 mm. long 5-8 mm. long

Wolffiella oblonga was first described in 1857 as a species of
Lemna by Philippi (7) from material collected near Santiago,
Chile. Hegelmaier (3) in 1868 in his classical monograph of the
Lemnaceae transferred the species to the genus Wolffia and erected
the subgenus Wolffiella to include it and several related species.
He expressed the belief that this subgenus should really be con-
sidered a distinct genus but was not prepared at that time to
make such a disposition of it. However, Hegelmaier (5) later
(1895) raised the subgenus Wolffiella to generic status and made
the combination Wolffiella oblonga ( Phil.) Hegelm.

The following material of Wolffiella oblonga (Phil.) Hegelm.
was studied. Chile: Santiago, May, 1857, Philippi (type collec-
tion, M). Argentina: Cordoba, May 9, 1898, Stuckert (M).
Uruguay: Montevideo, Pocitas, Herter 150: 70512 (M, UC).

Wolffiella oblonga (Phil.) Hegelm. has been reported from
California, an occurrence based upon two collections from near
San Bernardino, by Parish. A careful study of this material and
comparison with the type of W. oblonga eliminates that species

q

1938] MASON: FLOWERING OF WOLFFIELLA LINGULATA 24:7

from consideration in the California flora. These plants are
clearly small individuals of W. lingulata. It is a matter of inter-
est to note that a single specimen of the Parish collection shows
an immature flower in a very early stage of development.

Wolffiella lingulata was described as a Wolffia, subgenus Wolff-
ella, by Hegelmaier in 1868. To this in 1878 he added the vari-
ety W. lingulata var. minor. In 1898 Thompson (10) reduced this
variety to synonomy with W. oblonga ( Phil.) Hegelm. When the
subgenus Wolffella was raised to generic rank by Hegelmaier in
1896 the combination Wolffella lingulata (Hegelm.) Hegelm. was
made.

Wolffiella lingulata, as differentiated from W. oblonga by Hegel-
maier, was based largely upon size characters. His measure
ments for fronds of W. lingulata ranged from 4.7 to 6.1 milli-
meters in length and from 1.8 to 2.4 millimeters in width. For
W. oblonga he reported for the frond a length of 1.7 to 3 milli-
meters and a width of .6 to .85 millimeters. Later in describing
W. lingulata var. minor he noted that the size character between
the two species broke down, causing him to express doubt as to
the specific status of W. lingulata as distinct from W. oblonga.
Hegelmaier noted and figured the difference in the extent of the
air cavities in the fronds of the two species but made no particu-
lar point of this difference in his diagnosis. This character was
given emphasis, however, by J. D. Smith (8) and its importance
has been corroborated in the present investigation.

Thompson (9) largely on the basis of field and culture studies
of material collected near Bakersfield, California, and supple-
mented with specimen studies of material from Mexico and South
America, laid aside the doubts of Hegelmaier and rediagnosed
W. lingulata, calling attention to what appeared to him to be the
constant position of the stipe scar on the lower lip of the pouch
to the right of the median line of the frond. He stated that in
W. oblonga this scar occurs at the angle of the pouch to the right
of the frond. This, he maintained, consistently characterized the
small forms of W. lingulata as well as the large, while W. lingulata
var. minor on the other hand is in this respect identical with W.
oblonga and must be regarded as an elongated form of that spe-
cies. This character, he points out, serves as a ready means of
distinguishing these two species. The above observations of
Thompson’s relative to the position of the stipe scar have not
been substantiated in the investigations of the writer. As noted
above, both dextral and sinistral fronds have been observed and
the position of the stipe scar on the pouch is not a constant char-
acter. The dextral and sinistral position of the stipe in all proba-
bility is the result of genetic variation within the species and
probably only occurs as a result of sexual reproduction. In vege-
tative reproduction, as has been pointed out by Hegelmaier, the
stipe and costa of the daughter frond always develop on the side

248 MADRONO [Vol. 4

of the parent pouch opposite to the position of these organs on
the parent (pl. XX XV, figs. 7,11). Since the base of the daughter
frond faces the opposite direction from the base of the parent
frond, this necessitates that a dextral frond must produce dextral
offspring and likewise sinistral fronds must produce sinistral off-
spring. This could account for the uniformity of the plants re-
ported by Thompson from a sterile colony. They probably owed
their origin asexually to a single migrant individual. The posi-
tion of the stipe scar on the lower lip of the pouch or on its right
hand corner has been found to vary within a family colony, every
member of which descended asexually from a single individual.
Figures 9 and 10 of plate XXXV are mother and daughter
fronds. In figure 10 the stipe scar is on the lower margin of the
frond, whereas figure 9 illustrates an individual with the stipe
scar at the corner of the pouch. Figure 9 has the characters
attributed to W. oblonga in this respect. Since it can be demon-
strated that the size and shape of the frond is so dependent on
local and seasonal conditions the broad short character of the
frond attributed to W. lingulata by Thompson must be discarded
as a basis for differentiating the two species. Likewise the
“saber shaped” character attributed to W. oblonga by modern
writers does not hold. The great majority of the collections of
W. lingulata through most of the year are saber shaped. Figure
1 of plate XXXV is typical of W. lingulata according to the inter-
pretation of Thompson. It is a flowering specimen. Figures
2, 4, and 5 are of the saber shaped type. Flowers were found on
both types and no differences other than shape of the fronds
were noted between them. The differences between the two spe-
cies as observed by the writer (pl. XXXVI, figs. 1, 2) will per-
haps be best illustrated by a key.

Plants 1.5 to 4 mm. long; angle of vegetative pouch 40 to 50 de-

grees, the tip somewhat attenuate, the lips strongly rounded;

air chambers almost throughout the frond; pollen 11 to 15

microns in diameter, surface minutely wrinkled (fide Giar-

Cs LED i eA esr TETL HAA TOR CEM MEE MRC ARE NAAM AY be PL yo W. oblonga
Plants 4 to 9 mm. long; angle of vegetative pouch 60 to 90 de-

grees, the tip not attenuate; air chambers occupying not more

than half the frond; pollen 20 to 23 microns in diameter, the

surface muriculate 22 ib dunia eels ou ka acl ee eee W. lingulata

Since two species have been found flowering it would seem
that some information should now be forthcoming relative to the
generic status of Wolffella. Most writers have made their par-
ticular disposition of the group “pending the finding of flowers.”
In a plant so reduced in the structure of its vegetative and repro-
ductive parts it is scarcely to be expected that conclusive data
will be available to serve as a basis for descriptive differentiation.
Hegelmaier differentiated Wolfiella from Wolffia primarily on the
position of the axis of the vegetative shoot with respect to the
vegetative pouch. In Wolffia this shoot lies at the base of the

1938] MASON: FLOWERING OF WOLFFIELLA LINGULATA 249

IMMERSED

AREA

Prate XXXV. Wotrrietta tincutata (Hegelm.) Hegelm.
(See explanation of figures on page 250.)

250 MADRONO PWeles

reproductive pouch whereas in Wolffella it lies at or near one side
of the pouch. There is thus a tendency toward bilateral sym-
metry in Wolfia and toward asymmetry in Wolffiella. The dis-
covery of flowers adds nothing more conclusive than this vege-
tative character for separating the two genera. Although the
flowers vary in minor details that could scarcely have generic
significance, they are in all essential features alike.

University of California,

Berkeley, September, 1938.

LITERATURE CITED

1. GrarpveLti, M. L. Las flores de Wolffiella oblonga. Revista Argentina
de Agronomia 2: 17-20. 1935.

2.. GorseL, K. Zur Organographie der Lemnaceen. Flora 114: 278-305. 1921.

3. Hercetmairer, F. von. Die Lemnaceen, ein Monographische Untersuchung.
Leipzig. 1868.

A. Lemnaceae, in Martius, Flora Brasiliensis 32: 1-23. 1878.

5. ————————.. Systematische Ubersicht der Lemnaceen. -Engler’s Bot.
Jahrb. 21: 268-305. 1895.

6. Hicxs, L. E. Flower production in the Lemnaceae. Ohio Journ. Sci. 32:

115-131. 1932.

7. Purr, R. A. Plantarum novarum Chilensium, centuria quarta. Lin-
naea 29: 1-47. 1857.

8. Smiru, J. D. Wolffia (Wolffiella) gladiata Hegelm. Bull. Torrey Bot.
Club 7: 64-65. 1880.

9. THompson, C. H. The ligulate Wolffias of the United States. Missouri
Bot. Gard., 7th Ann. Rept. 101-111. 1896.

10. ————————. Revision of the American Lemnaceae occurring north of
Mexico. Missouri Bot. Gard., 9th Ann. Rept. 1-42. 1898.

EXPLANATION OF THE FIGURES

Puate XXXV. Wolffiella lingulata (Hegelm.) Hegelm. Fig. 1. Ligulate
type of frond showing floral cavity with pistillate flower in anthesis. Fig. 2.
Frond showing position of flower with respect to immersed area of plant. Fig. 3.
Profile view of staminate flower showing its position relative to water surface.
Figs. 4,5. Family colonies of saber shaped fronds. Fig. 6. Upper epidermis
of flowering frond showing pigment cells and stoma. Fig. 7. Parent and daugh-
ter frond showing position of daughter stipe and pouch with respect to the
parent. Fig. 8. Stipe of frond of the colony stage. Note abscission layer.
Figs. 9,10. Daughter and parent fronds showing variation in position of stipe
scar in a single family. Fig. 11. Separation of parent and daughter frond.

Prate XXXVI. Fig. 1. Frond of Wolfiella oblonga from type collection.
Note extent of air chambers in tissue of frond. Fig. 2. Frond of Woffella
lingulata. Note extent of air chambers. Figs. 3-18. Wolffella lingulata.
Fig. 3. Mature daughter frond still attached to parent. Note flower bud and
reduced size of flowering frond as compared to size of parent. Figs. 4-8. Stages
in flowering and fruiting: 4, pistillate flower in anthesis, staminate flower not yet
emerged; 5, emergence of staminate flower; 6, dehiscence of stamen; 7, degen-
eration of stamen and development of fruit; 8, mature fruit. Fig. 9. Mature
fruit with persistent style. Fig.10. Pollen grains, x 500. Fig.11. Two seeds,
left hand one showing cavity in which lies the operculum of the testa. Fig. 12.
Mature fruit in dead parent frond. Figs. 13-18. Stages in development of
seedling. Terminology of fig. 17 adapted from Goebel (2).

1938] MASON: FLOWERING OF WOLFFIELLA LINGULATA 251

) .

GLOBULE OF
STIGMATIC
STAMINATE SECRETION
FLOWER

PISTILLATE
FLOWER

STAMINATE. PISTILLATE
FLOWER FLOWER 4.
AFTER ANTHESIS

Sy
STAMINATE

FLOWER PISTILLATE

AFTER FLOWER
ANTHES!S

AFTER ANTHESIS

YOUNG FRUIT
\7 DAUGHTER
FROND
COTYLEDONARY
¥ SHEATH
U TESTA
SrAne ( OPERCULUM
MATURE HYPOCOTYL
FRUIT PERSISTANT y

STYLE

Ha
yk PARENT
SOAs FROND
ee

Pirate XXXVI. Wourrietta. (See explanation of figures on
page 250.)

252 MADRONO [Vol. 4

THE WESTERN AMERICAN SPECIES OF PAEONIA

G. Lepyarp STessIns, JR.

In all recent treatments of the genus Paeonia in the western
United States, only one species has been recognized, P. Brownii
Dougl. The range of this “‘species,’ however, is most unusual.
Paeonia occurs commonly in cismontane southern California at
low altitudes. It is rare in central California, but reappears as
a frequent or common plant at medium altitudes, in the northern
part of the North Coast Ranges and in the northern Sierra
Nevada, and from there extends northward to British Columbia
and eastward to Wyoming.

Struck by this remarkable discontinuity in range, Pe
systematists have maintained that the peony of southern and
south central California is specifically distinct from that of the
northwest. Nuttall, whose manuscript description is quoted by
Torrey and Gray (8), described the southern form as Paeonia
californica, which, he stated could be distinguished from the
northern P. Browni by its “smaller, less divided leaves, which
are deep green on both sides, and the leaflets bifid or trifid, never
pinnatifid,’ as well as by possessing three rather than five carpels.
Nuttall’s type of P. californica was collected near Santa Barbara;
Douglas’ type of P. Brownii in Oregon, “near the confines of per-
petual snow on the alpine range of Mt. Hood.’ Unfortunately,
additional material of both species revealed that, with the excep-
tion of green rather than glaucous leaves, all of the character-
istics mentioned by Nuttall occur also in the northern P. Brownit.
Many plants of this species have leaves smaller than those of
typical P. californica; in some specimens of P. Brownii the leaves
are not more dissected than are those of the southern plant; the
number of carpels varies from two to five in both. Probably
recognizing these facts, Gray (2) as well as Brewer and Watson
(1) considered P. californica a synonym of P. Brownii. The
authors of recent California floras, Jepson (4, p. 873) and Munz
(6, p. 170) have also recognized only P. Brownii, and have not
even mentioned P. californica as a synonym.

The strongest case for recognizing Paeonia californica as a
distinct species was presented by E. L. Greene (3). He drew
a graphic picture of the ecological differences separating the two
but failed to give any valid diagnostic characters to distinguish
between them, except leaf color and shape of leaf segments.
Using these characteristics, Lynch (5) recognized P. californica
as a variety of P. Brownit.

The writer became interested in this species problem through
cytological and systematic studies of the Old World species of the
genus. Upon examining herbarium specimens, it was found that
after a little study one could tell almost at a glance, without look-
ing at the label, whether a specimen was from southern California

1938] STEBBINS: PAEONIA 253

»

Taste 1.—Comparison of Paeonia Brownii and P. californica

P. Brownu P. californica

Hasir
Number of flowering

stems per plant 1-16 5-30
Height of flowering

stems 2-4 dm. 3.5—7.5 dm.
Number of leaves per

stem 5-8 (av. 6-7) 7-12 (av. 9-10)

Character of branching stems usually simple,
sometimes with 1 or
2 short branches

stems always branched
or with rudiments of
branches in axils of up-
per leaves; in luxuriant
plants the larger primary
branches bear secondary
branches

CATAPHYLLS

Apex obtuse acute

LEAVES

Color glaucous green

Texture thick, usually some- thin, easily wilting

what fleshy

Base of primary abruptly contracted, cuneate, sessile or with

segments petiolulate short, alate petiolules

Terminal secondary seg-
ment of middle cauline
leaf, length 3.5—-6.0 cm. 2.8—7.5 cm.

Terminal secondary seg-
ment of middle cauline

leaf, width 2.5—-5.5 cm. 1.1—4.5 em.
Ratio of length to
width 0.8-1.4 1.5-3.0

Number of ultimate
lobes on terminal pri-
mary segment of mid-

dle cauline leaf 13-44 10-17
Shape of ultimate lobes elliptic, usually lanceolate or narrowly
obtuse elliptic, usually acute
FLOWERS
Petals rotund to orbicular elliptic
Length of largest petal 8-13 mm. (av. 9-11) 15-25 mm. (av. 19-21)
Width of largest petal 10-15 mm. 11-18 mm.
Ratio of length to width 0.7-1.0 1.1-1.6
Petal, color of center maroon or bronze deep blackish red
Petal, color of margin yellowish or greenish pink (sometimes a narrow
yellowish margin at the
apex)
Length of longest fila-
ments 4,.2-5.5 mm. 6.0-8.5 mm.
Length of longest
anthers 3.3-5 mm. 3.5-7 mm.

or from the north. The exact differences between the two, how-

ever, were not easily defined, largely because the leaves of
Paeonia are rarely well spread out in the preparation of speci-

254 MADRONO [Vol. 4

mens, the bulky flowers are usually crushed in such a manner that
little can be told of their fresh condition, and there are rarely
enough flowers on a specimen to permit dissection.

For this reason, the writer has paid particular attention to
Paeonia on collecting trips. Several collections of P. Brownii
were made during the seasons of 1936 and 1987 in Nevada,
Sierra, Lassen, and Shasta counties in northeastern California,
and a small colony was discovered on the northeast side of Mount
Hamilton, Santa Clara County, in the Coast Range. This is a
southerly extension of the known range of typical P. Brownii;
the nearest known locality to the northward is in Lake County,
about 150 miles distant by air line. The southern plant was
studied and collected in 1937 at the type locality near Santa
Barbara, as well as near San Luis Obispo and at the foot of Cajon
Pass, San Bernardino County; in 1988 its most northerly known
locality, Pine Canyon, west of King City, Monterey County, was
visited. In making each collection, a number of flowers were dis-
sected before pressing as described elsewhere (Stebbins, 7).

On the basis of these observations, the existence of a number
of real distinctions between P. Brownii and P. californica was
established. These are stated in Table 1 and the most important
summarized in the following key:

Stems 2-4 dm. high, bearing 5-8 leaves; leaves more or less

glaucous, their primary divisions abruptly’ contracted at

the base to distinct, often elongate petiolules, the ulti-

mate lobes elliptic; petals rotund or orbicular, mostly

broader than long, the larger 0.8-1.3 cm. long when fully

spread out, definitely shorter than the inner sepals ..... 1. P. Brownit
Stems 3.5-7.5 dm. high, bearing 7-12 leaves; leaves green, not

glaucous, their primary divisions cuneate at the base,

often sessile, the ultimate lobes lanceolate or narrowly

elliptic; petals elliptic, longer than broad when fully

spread out, the larger 1.5-2.5 cm. long, slightly longer

than: the: inner ‘sepals. 32 cc tins. re ee tee eee 2. P. californica

1. Partonta Brownnt Dougl. ex Hook. Fl. Bor. Am. 1: 27.
1829; Torr. & Gray, Fl. N. Am. 1: 41: (1838, | Dlustrations:
Bot. Reg. 25, t. 830. 1889; Journ. Royal Hort. Soc. Lond. 12:
433. fig. 24,1890. (pl. XXXVII, figs. f-n.)

Northern California, from Santa Clara and Tuolumne coun-
ties north to British Columbia, east to Wyoming. In Cali-
fornia mostly at altitudes of 3000 to 6000 feet. (900 to 1800
meters). The following specimens in the herbarium of the Uni-
versity of California (UC) and in the Dudley Herbarium, Stan-
ford University (DS), are typical. Ipano: Silver City, Owyhee
County, Macbride 943 (UC, DS). Wasuineron: Swauk Creek,
Wenatchee Mts., Quick 1042, 1054 (UC). Oregon: Rock Creek,
Morrow County, alt. 1040 m., Leiberg 95 (UC). Catirornia:
Quartz Valley, Siskiyou County, Butter 1229 (UC, DS); Jame-
son Creek, Plumas County, Hall 9307 (UC); Bartlett Mt., Lake

1938] STEBBINS: PAEONIA 255

County, Abrams 12416 (DS); Mount Hamilton, Santa Clara
County, H. K. Sharsmith 3947 (UC).

The exact eastern limits of Paeonia Brownii are uncertain.
Rydberg (Rocky Mt. Flora, p. 315) includes Utah in the range;
Tidestrom (Fl. Utah and Nevada, p. 202) questions its occurrence
there but includes Alberta, which Rydberg does not. In answer
to requests by the writer, Dr. Ellsworth P. Killip of the United
States National Herbarium (US), Mr. C. A. Weatherby of Gray
Herbarium (G) and Dr. H. A. Gleason of the New York Botanical
Garden (NY), have examined the material of this species in their
collections and report no specimens of P. Brownii from either
Utah or Alberta. One collection is reported from Wyoming:
Jackson’s Hole, Lincoln County, Payson & Payson 2196 (G, NY).

2. Pakonia cCALirorNica Nutt. ex Torr. & Gray, Fl. N. Am. 1:
Ain) 838. Greene, Garden and Forest 3::356. . 1890. .)(P-
Browni var. californica Lynch. Journ. Royal Hort. Soc. Lond. 12:
433. 1890). Illustrations: Lynch. l.c. fig. 25, leaf; Parsons,
Wild Fl. Calif. 841. 1897; Davidson, California Plants in their
Homes, fig. 31, 1898; Armstrong, Field Book West. Wild FI. 139.
1915; Pickwell et al., Spring Wild Flowers of the Open Field,
Western Nature Study 2: fig. 81. 1981; Munz, Man. S. Cal.
Bot. 170, fig. 76, 1935; Thurston, Wild Fl. So. Cal. 326, fig. 511,
1936. (pl. XXXVII, figs. a—e.)

Southern and south central California, from Monterey to San
Diego County, from sea level to 4000 feet (1212 meters) (fide
Munz, 1935). The following specimens are typical. Cati-
FORNIA: east of Bryson, Monterey County, Ferris 8450 (UC, DS) ;
Santa Ynez Mts., Santa Barbara County, Elmer 8786 (UC, DS);
Santa Monica Mts., Los Angeles County, alt. 425 m., Clokey &
Templeton 4447 (UC); San Bernardino Valley, San Bernardino
County, alt. 360—400 m., Parish 6820, 11713 (UC) ; Encinitas, San
Diego County, Brandegee in 1884 (UC).

As will be seen from the table, most of the differences be-
tween the two species are associated with the greater luxuriance
of growth in Paeonia californica. This species is larger and better
developed in all of its parts than P. Brownii, except that no sig-
nificant differences between the two could be found in the outer
sepals, the carpels at anthesis, the mature follicles, and the seeds.
Paeonia californica, judging from its range, is not hardy, and is a
mesophyte. Although the climate of its habitat is semiarid so
far as the season as a whole is concerned, P. californica completes
its active growth during the rainy season, when the ground is
more or less moist, and lies dormant during the dry season, de-
pending on food stored in its thick roots. Its leaves and stems
wilt very readily.

Paeonia Brownii, on the other hand, is definitely hardy, since
its shoots sometimes push their way up through banks of snow.
It is also semi-xerophytic; at least the latter part of its seasonal
growth takes place after precipitation has almost or entirely

256 MADRONO [Vol. 4

ceased, and the ground has become quite dry. Its fleshy leaves
retain their turgor for many hours after the stem is broken.

On the basis of these facts, the question arises as to whether
or not the differences between Paeonia Browniu and P. californica
are due to the direct stimulus of the environment, and whether
they could be transformed one into the other by transplanting.
The critical experiments to test this point have not yet been per-
formed, but all available evidence indicates that such transforma-
tion would not take place. Many experimenters, notably Tures-
son (9), have shown that the large majority of adaptations to
different habitats within the same or closely related species are
genetically conditioned.

Within each of these two species of Paeonia variation in the
characters by which they differ from each other is not noticeably
correlated with differences in habitat. Plants of P. californica in
Monterey County, central California, are practically indistin-
guishable from those of the same species occurring about San
Diego, 350 miles farther south, which receive about half as much
rainfall. Those of P. Brownitt growing in central California, on
Mount Hamilton, are very similar to those found in the We-
natchee Mountains of Washington, 750 miles to the north.
Furthermore, P. Brownit shows evident “conservatism” in the
southern part of its range. It is rare in the Coast Ranges of
California and is apparently retreating with the advent of a
warmer climate. Many of the plants in the Mount Hamilton
colony, the southernmost known locality, produce only small,
abortive buds and no flowers, an indication that the species is
there dying out. All evidence points to the assumption that
plants of P. Browni are not only unable to adapt themselves to a
warmer climate, but in addition that they cannot, either by segre-
gation or by mutation, produce offspring that can tolerate a
warmer climate. Paeonia californica, on the other hand, shows
every sign of advancing with the increasing warmth of the cli-
mate. It is a rather common plant in suitable spots up to the
northern limit of its range, and at its northernmost known station
is abundant and very luxuriant. It has not, however, been able
to colonize the localities within its range which have a climate
corresponding to that required by P. Browni. In the higher
mountains of southern California, which have a climate and flora
similar to that of the northern Sierra Nevada where P. Browni
is abundant, there is no Paeonia at all. At present, therefore, all
evidence warrants the assumption that neither by physiological
nor by genetical change can either of these species become trans-
formed into the other.

Another possible explanation is that the two forms represent
geographic variations—varieties, subspecies, or ecotypes (Tures-
son, 10) of a single species. The crucial test of this, hybridiza-
tion of the two, would be a long-time proposition since most
species of Paeonia do not flower until they are four or five years

1938] STEBBINS: PAEONIA 257

old. Furthermore, the significance of hybridization tests would
be complicated by the fact that both species normally have 60 to
70 per cent of pollen sterility, due to their anomalous cytological
condition (Stebbins and Ellerton, in press). At present, how-
ever, the facts that no intermediates exist and that hybrids could
not be produced except under highly artificial conditions of culti-
vation, speak strongly in favor of maintaining Paeonia Browniu
and P. californica as distinct species.

Paeonia californica is a relatively constant species. Within
P. Brownii, on the other hand, there is a considerable range of
variation, particularly in the amount of dissection of the leaves
and in the shape of the segments. In the northern Sierra Nevada
a form occurs with leaves much dissected, and frequently with
smaller flowers than are typical for the species. Since, how-
ever, plants with leaves and flowers of the normal type occur
within the range of this Sierran variant, the writer does not con-
sider it well enough marked for recognition as a distinct variety
or subspecies.

The phylogenetic relationship between the two species is best
determined by comparing them with the Old World species of
Paeonia. Paeonia Brownii and P. californica constitute a distinct
subgenus, Onaepia (Lynch, 5), which is characterized by small
petals; a very prominent disk, usually divided into separate seg-
ments; and cylindrical rather than ovoid seeds. The Eurasian
species are segregated into two subgenera, Moutan comprising the
shrubby, and Paeon, the herbaceous species. In spite of the
shrubby habit, Moutan is nearer to Onaepia than is Paeon. The
“tree peonies” of China have, like the American species, a very
prominent disk, and both have large seeds with dull coarsely
rugose surfaces; the seeds of Paeon are smaller, relatively smooth,
and often shining. Furthermore, one of the two species of sub-
genus Moutan, P. Delavayi Franch. of southwestern China, has
petals almost as small as those of P. californica, and in the typical
form of P. Delavayi they are of a similar reddish color. Paeonia
Delavayi resembles P. californica also in its sepals and carpels and
in the shape of its leaves, although those of the Chinese species
are much larger and more dissected. In some specimens of P.
californica the disk segments are relatively thin and are partly
fused with each other, recalling the conspicuous, continuous disk
of P. Delavayi. Both species are characterized by rather indefi-
nite branching, the branches generally bearing well developed
leaves as well as a terminal flower bud. In all other species of
Paeonia (except occasional luxuriant individuals of P. Brownii),
the side branches bear only a terminal bud, with sometimes a
reduced leaf below it. Since P. Delavayi must be considered the
most primitive species of Paeonia extant, (Stebbins, 7) the line
which gave rise to P. californica and P. Browni must have
branched off from the rest of the genus very early in its history.

258 MADRONO [Vol. 4

The fact that the closest connection between the Old and the New
World species of Paeonia is through species endemic to such re-
mote and different regions as southern California and south-
western China suggests that this connection is very ancient. The
Old World species, P. albiflora Pall. of Manchuria, from which
the well known cultivated peony is derived, is geographically
nearest to P. Brownii and occupies a somewhat similar habitat,
but it is obviously very remotely related to both P. Brownw and
P. californica. In all of the most important morphological charac-
ters by which P. Browniu differs from P. californica, it diverges
also from all of the Old World species. In shape and texture of
leaves, shape and color of petals, and in habit P. Brownii is quite
unique. It represents, therefore, an offshoot which has diverged
more in morphological characters from the common ancestor of
the American peonies than has P. californica. The latter species,
however, shows more ecological divergence from other species
of the genus, since it is the only peony that is not frost-hardy.
The most likely hypothesis as to the course of evolution of the
two species is that their common ancestor was mesophytic and
frost-hardy, as are all Old World species, and resembled mor-
phologically P. Delavayi and P. californica. This ancestor prob-
ably was present in California in Mesozoic or early Tertiary time,
and gave rise to a considerable number of ecological and mor-
phological types. Of these all that were mesophytic and frost-
hardy perished when the climate of California became warmer
and more arid. Paeonia californica persisted in southern Cali-
fornia by evolving a change in its period of dormancy from the
cold winter to the dry summer, but retained most of the mor-
phological characters of its original ancestor. Paeonia Brownii,
on the other hand, retained its frost-hardiness and winter period
of dormancy, but became adapted to dry summers by evolving
a series of morphological specializations. Both species, there-
fore, are derived types, but morphologically P. californica is
undoubtedly the more primitive.

Division of Genetics,
University of California, Berkeley, April, 1938.

EXPLANATION OF THE Figures. PLate XXXVII

Figs. a-e, Paeonia californica (Santa Ynez Valley, California, Stebbins
2090): a, terminal primary segment of median cauline leaf, xX %; 6, the three
innermost sepals, X 1; c, the largest and smallest petals, xX 1; d, two stamens, <1;
e, disk and follicles at anthesis, X< 1.

Figs. f-i, Paeonia Brownii (Wenatchee Mts., Washington, Quick 1042):
f, terminal primary segment of median cauline leaf, X 4%; g, the largest petal,
X1; h, two stamens, X 1; i, disk and follicles at anthesis, x 1.

Figs. j-n, Paeonia Brownii (Sierra County, California, Stebbins & Jenkins
2157): j, terminal primary segment of median cauline leaf, x 42; k, the five inner-
most sepals, X 1; 1, the largest and smallest petal, x 1; m, two stamens, <1; n, disk
and follicles, x 1.

1938] STEBBINS: PAEONIA 259

Prate XXXVII. Western AMERICAN SPECIES OF PAEONIA.

260 MADRONO [Vol. 4

LirerATURE CITED

1. Brewer, W. H., and S. Watson. Paeonia L., in Botany of California 1: 13.
1876.

Gray, A. Paeonia L., in Synoptic Flora of North America 1(1): 56. 1875.

GREENE, E. L. The California Paeonias. Garden and Forest 3: 356. 1896.

Jepson, W. L. A Manual of the Flowering Plants of California. 1925.

Lyncn, R. I. A New Classification of the Genus Paeonia. Journ. Royal
Hort. Soc. Lond. 12: 428-445. 1890.

Munz, P. A. Manual of Southern California Botany. 1935.

Stessins, G. L., Jr. Notes on the systematic relationships of the Old World
species and of some horticultural forms of the genus Paeonia. Univ.
Calif. Publ. Bot. (in press).

8. Torrey, J.. and A. Gray. Paeonia L., in Flora of North America 1: 41.
1838.
9. Turesson, G. The plant species in relation to habitat and climate. Hered-
itas 6: 147-236. 1925.
10. ———————.. Zur Natur und Begrenzung der Arteinheiten. Hereditas
12: 324-334. 1929.

emer

HANGING GARDENS OF THE CANARY ISLAND
DATE PALM

Ira L. Wiccins

The practice of trimming the older leaves from the trunks of
palms used in ornamental plantings provides a temporary arbo-
real habitat for a number of plants that normally grow only on
the ground. The row of Canary Island date palms on either side
of Palm Drive on the campus of Stanford University, California,
annually supports a total population of many hundred individual
plants and a surprisingly large number of species. The relation-
ships between these plants that are normally ground-dwellers
and their arboreal supporters, the fluctuations in the “hanging
gardens” from year to year, and the variations in the tenacity of
such wanderers marooned above their normal sphere are sources
of considerable interest.

Many palms growing under wild conditions retain their leaves
over a period of years, thus forming a close thatch that shades
out any seedlings that may, upon rare occasions, begin growth
on their trunks. But even in such trees, the functional leaves,
extending upward at sharp angles, provide catchment basins in
the bases of their petioles for decaying organic matter and a
small quantity of dust. In cultivated trees a considerable por-
tion of this material remains in the fissures between the basal
parts of the petioles after the bulk of the leaves has been pruned
from the trunk. The functional leaves above form a natural
drainage system that directs much of the intercepted rainwater
into the basal parts of the petioles. Some of this water is re-
tained in the cups formed by the petiolar bases, some of it trickles
over the rims to enter those below and fill the lower reservoirs.
Thus, during the course of a heavy rain the entire trunk of one

1938] WIGGINS: HANGING GARDENS 261

of these date palms may become a series of miniature reservoirs,
each containing a small quantity of decayed organic matter and
rainwater. ;

Seeds lodged in these pockets find conditions favorable for
germination and rapid growth, with the result that many of the
palms become decorated with various annual and perennial seed-
lings. Many of the herbaceous annuals contributing to these
hanging gardens live through their entire life cycles before the
water supply fails. Such plants reseed the petiolar cups for the
next season’s growth so the gardens flourish year after year.
Even the perennials survive for a surprisingly long time, though
few, if any, live long enough in this habitat of limited supplies
to produce fruit. Ferns furnish an exception among the peren-
nials, for they produce fertile fronds regularly.

Casual observations made over a period of ten years or more
indicate that there are four principal agencies that are effective
in sowing seeds in the petiolar bases of the palms. They are: 1,
gravity; 2, wind; 3, animal and bird carriers; 4, direct deposition
of seed by annuals growing against the trunks of the trees.

Gravity is responsible for the presence of numerous specimens
of young palms in the petiolar cups of the parent trees. Falling
fruits lodge among the leaf-bases in great numbers and some of
their seeds grow. Occasionally seeds of Acacia and acorns from
the oaks fall among the leaves of the palms. Although Eucalyp-
tus trees tower above the palms at a number of places and some
of the seeds shattering from the capsules must lodge in the
petiolar cups, no seedlings of Eucalyptus appear in the arboreal
habitat. Their absence is doubtless to be attributed to some fac-
tor in the petiolar gardens that is unfavorable to the germination
of the seeds of Eucalyptus rather than to the failure of the force
of gravity!

Birds and small rodents carry acorns, pine nuts, seeds of some
species and berries of others from the parent plants to the leaf-
bases of the palms before the outer parts of the leaves are re-
moved, or, in the cases of woodpeckers, jays, and squirrels, may
hide seeds and nuts among the petiolar bases some distance below
the leaf-crown. Robins drop fruit of Photinia, Pyracantha, and
Cotoneaster among the leaves. Waxwings carry these fruits, as
well as those of Schinus molle, from fruiting plants to other lo-
calities, and the purple house finch after eating the berries of
Sambucus, frequently takes refuge in the palm trees. The habit

_of wiping beaks on the perches doubtless results in the deposition
of some seeds in the crevices among the petiolar bases.

The wind carries seeds of such species as Ailanthus glandulosa
and Sonchus oleraceus to the arboreal habitats, and the same
agency is responsible for the presence of Polypodium and Dryop-
teris on the sides of palm trunks high above one’s head. It is

262 MADRONO [Vol. 4

dificult to explain the absence of such seedlings as those of
Ulmus, Platanus, Senecio and others with wind-borne seeds.

The most interesting group, however, is made up of those
species that do not have wind-blown seeds, do not seem to be
used extensively as food by birds and animals, and grow on the
ground below the trunks on which some of their number appear
season after season. Periodic observations have shown that such
annuals as Stellaria media, Anagallis arvensis, Festuca megalura,
Silene gallica, Montia perfoliata, and Bromus rigidus slowly ciimb
higher and higher from one year to another until they occur
from the ground to the leaf-crown. An occasional plant of one
of these weedy species missed by the gardener’s hoe grows against
the base of the tree. Its seed capsules or fruiting spikes drop a
few seeds into the crevices between the petiolar bases a short dis-
tance above the soil and the plant is on its way upward. The
plants resulting from the seeds thus sown also grow in close con-
tact with the tree-trunk, mature fruit, and deposit more seeds a
few inches higher up the trunk of the supporting palm tree.
This progressive ascent may account for the presence of an
almost solid growth of annuals from the ground to the leaf-crown
on some trees, the almost total absence of them on other trees
in the same row.

There is a noticeable lack of hanging gardens on the trunks
of fan palms. The attachment of the petioles to the trunk of
Washingtonia filifera, for example, is such that water is not held
in the bases of the petioles. The basal part of the petiole bifur-
cates, each branch running diagonally downward and about one
fifth of the way around the trunk before the main vascular supply
enters the trunk of the tree. The increase in diameter of the
trunk after the initial departure of the young leaf-traces further
splits the base of the petiole and tears the tissue between many
of the smaller vascular strands running into the trunk. Thus a
considerable portion of the petiolar base is unattached to the
trunk of the tree, so water running down from the leaves above
percolates through the fibrous strands enwrapping the trunk to
enter the soil at the base of the tree. Therefore the trunk of the
tree and the crevices between the petiole bases dry out too
rapidly to permit the growth of even the short lived annuals.

Some perennials survive in the crevices among the petiolar
bases on the date palms for several years. Seedlings of Pinus
radiata five years old, of Pinus Sabiniana three years old, and of
Quercus agrifolia nine years of age are not infrequent on the
trunks of date palms along the drive on the Stanford campus.
Plants of Polypodium californicum and of Dryopteris arguta are now
under observation for the sixth successive year. A specimen of
Eriobotrya japonica (loquat) that started as a seedling in a
petiolar base of a date palm in front of the Museum Building
reached the leaf-crown of the palm, nearly twenty feet above its

1938] MUENSCHER: FLORA OF MOUNT BAKER 263

roots, before being removed by the gardeners. However, this
seedling started only a couple of feet above the ground, so its
roots soon found their way into the soil, and its period of depen-
dence on the scanty supply of soil and moisture in the petiolar
cups was comparatively short.

It would be interesting to know something about the fluctua-
tions in the available moisture in these cups, and whether the
perennials die on account of water shortage or through the
exhaustion of the essential mineral elements entrapped in the
petiolar crevices.

The following list includes only those species observed grow-
ing on the date palms on the Stanford campus and in park ways
near my home. A more extensive survey might expand this list
considerably.

Acacia retinodes Schlecht.

Ailanthus glandulosa Desf.

Anagallis arvensis L.
Avena fatua L.
Bromus mollis L.

Montia perfoliata (Donn)
Howell

Phoenix canariensis Hort.

Pinus radiata Don

Pinus Sabiniana Dougl.

Prunus communis L.

Prunus ilicifolia Walp.

Bromus rigidus Roth.
Cupressus Macnabiana

Murr. Polypodium californicum
Cupressus macrocarpa Kaulf.
Hartw. Quercus agrifolia Née

Dryopteris arguta (Kaulf.) Pyracantha sp.

Watt. Sambucus glauca Nutt.
Eriobotrya japonica Lindl. Silene gallica L.
Echinocystis fabacea Naud. Solanum nigrum L.
Festuca megalura Nutt. Sonchus oleraceus L.
Festuca myuros L. Stellaria media (L.) Cyr.
Hordeum murinum L. Thuya orientalis L.

Verbena prostrata R. Br.

Dudley Herbarium,
Stanford University, July, 1938.

ADDITIONS TO OUR KNOWLEDGE OF THE FLORA OF
MOUNT BAKER, WASHINGTON

W. C. MvuENSCHER

Mount Baker, with its cap of perpetual ice and snow, attain-
ing a height of 10,750 feet above sea level, dominates the land-
scape of Whatcom County, Washington, the northwest corner of
the United States. In 1929 St. John and Hardin (3) published
a flora of this area reporting its known flora as consisting of 334
species and varieties.

264 MADRONO [Vol. 4

As a result of the writer's botanical explorations of Mount
Baker, made between 1910 and 1937, its flora is well represented
in the herbarium of Cornell University. The present list of
vascular plants consisting of additions to the known flora of the
Mount Baker region, as delimited by St. John and Hardin, is
largely based upon the writer's observations and collections. A
record of the occurrence of a few of these species has been pub-
lished previously (1,2). Mr. J. W. Thompson of Seattle, Wash-
ington, has published records of seven species (4, 5, 6) and has
communicated recently by letter the occurrence of 18 additional
species. With the exception of Mr. Thompson’s records, the
present list is based entirely upon collections deposited in the
herbarium of Cornell University. Introduced as well as native
species are included.

The present list increases the known plants of the region, as
published by St. John and Hardin, by 13 families, 90 genera and
228 species, an increase by 68 per cent of the previously reported
flora. At least forty weeds occur, most of which have been intro-
duced in openings along the lower trails and few roads that pene-
trate the area. The presence of so many introduced species indi-
cates that the flora of the region, although still mostly in a
primitive state, is no longer as isolated as formerly, even though
it is located in a national forest.

Acknowledgment is due to Professor K. M. Wiegand for assis-
tance in the determinations in several critical genera and to Dr.
R. T. Clausen who determined the species of Botrychium.

POLYPODIACEAE

AsPLENIUM TricHomaNneEs L. Rocky slopes in lower forest.

ASPLENIUM VIRIDE L. Moist ledges near timber line.

CHEILANTHES GRACILLIMA D.C. Eaton. Mt. Hermann, J. W. Thompson.

CRYPTOGRAMMA DENSA (Brack.) Diels. Rocky ledges near timber line.

Dryopteris Puecorrerts (L.) C. Chr. At base of Panorama Dome, J. W.
Thompson.

PotysticHum Lemmonu Underw. Rocky ledges above timber line.

Preripium aquitinum (L.) Kuhn. var. Ltanucinosum (Bong.) Fern. Common
on lower slopes.

Woopnsta scoputina D. C. Eaton. Mt. Hermann, J. W. Thompson.

OPpHIOGLOSSACEAE

BorrycHIuM LANCEOLATUM (Gmel.) Angstr. Alpine meadows.

Borrycuium Lunaria (L.) Swartz. Alpine meadows.

BotrycHIUM MULTIFIDUM (Gmel.) Rupr. subsp. strairotium (Presl) Claus. Moist
banks about mountain meadows.

EQUISETACEAE

E@vuIsETUM HYEMALE L. Springy slopes in lower forest.
EevuiseTumM timosum L. Marshy banks of streams and ponds.

LYCOPODIACEAE

LycoropIuM ANNOTINUM L. Moist slopes in the lower forest.
Lycoropium ogscurum L. Dry exposed ridge, Sulphur Creek.
Lycoropium Setaco L. Among mosses in lower forest.

1938] MUENSCHER: FLORA OF MOUNT BAKER 265

SELAGINELLACEAE
SELAGINELLA Wattace! Fier. Dry rocky slopes near timber line.

TsOETACEAE

IsorTEs ECHINOSPoRA Dur. var. Braun (Dur.) Englm. In shallow water, Baker
Lake.

PINACEAE

PicEA SITCHENSIS (Bong.) Traut. and Mey. Infrequent in lower forest along
Baker River and Swift Creek.
Pinus contorta Dougl. Dry burned over slope near Sulphur Creek.

TYPHACEAE
TypuHa Latirot1a L. Marshy shores of Baker Lake.

SPARGANIACEAE
SPARGANIUM MULTIPEDUNCULATUM (Morong) Rydb. Baker Lake; Baker River.

PoTAMOGETONACEAE

PoTAMOGETON AMERICANUS Cham. and Schlecht. Baker River near Baker Lake.
POTAMOGETON EPIHYDRUsS Raf. Baker Lake; Baker River.
POTAMOGETON TENUIFOLIUS Raf. Baker Lake; Baker River.

GRAMINEAE

AGROPYRON TRACHYCAULUM (Link) Steud. Dry meadows, Skyline Ridge.

Acrostis THURBERIANA Hitche. Alpine meadows, J. W. Thompson.

AIRA CARYOPHYLLEA L. Common on dry soil in openings in the lower forests.

Bromus ciziatus L. Moist banks.

Bromus TEctorum L. A weed about Glacier.

CALAMAGROSTIS CANADENSIS (Michx.) Beauv. var. scasra (Presl) Hitche. Wet
meadows.

DactTyis GLoMERATA L. Introduced along roads, J. W. Thompson.

Etymus ciaucus Buckl. Meadow on Skyline Ridge.

Festuca rusprA L. Dry slopes.

FEstTuca suBULATA Trin. Baker Lake region.

Festuca supina Schur. Alpine meadows.

Guyceria ELATA (Nash) Hitche. Swampy ground along lower streams.

Hotcus tanatus L. A weed along the lower roads and trails.

MUHLENBERGIA FILIFORMIs (Thurb.) Rydb. Near Austin Pass, J. W. Thompson.

PHLEUM PRATENSE L. Introduced along roads and trails, J. W. Thompson.

Poa atpIna L. On ledges in alpine meadows.

Poa annua L. Baker Lake region.

Poa arctica R. Br. Heliotrope Ridge, J. W. Thompson.

Poa Epiuis Scribn. Among rocky ledges, Skyline Ridge.

Poa GRACILLIMA Vasey. Common in alpine meadows.

Poa patustris L. Introduced along roads, J. W. Thompson.

Poa pratensis L. Introduced along roads, J. W. Thompson.

Poa sEcunpDA Presl. Rocky alpine slopes, J. W. Thompson.

TRISETUM CERNUUM Trin. In moist forests.

CYPERACEAE

Carex ABLATA Bailey. Wet meadows.

Carex arcta Boott. Open areas in the lower forest.

CarEX FESTIVELLA Mack. Marshes and wet meadows.

Carex LAEVICULMIS Meinsh. Wet meadows and along streams.

CAREX LEPORINELLA Mack. Alpine meadow, Heliotrope Ridge.

Carex LEPTOPEDA Mack. Marshy places in lower forests.

Carex Pirert Mack. Grassy slopes.

Carex Rossit Boott. Meadows near timberline, J. W. Thompson.

Carex ROsTRATA Stokes var. uTRICULATA (Boott) Bailey. In shallow water, Baker
Lake. }

» ie

266 MADRONO [Vol. 4

CarREX sITCHENSIS Bailey. Sloughs and wet banks of lower streams.

CaREX VEsICARIA L. var. Magor Boott. Marshes about Baker Lake.

DvuLICHIUM ARUNDINACEUM (L.) Britt. Marshes about Baker Lake.
ELEocHaris optusa (Willd.) Schultes. Meadow near Glacier, J. W. Thompson.
Scirpus microcarPpus Presl. Common in wet places in the lower forest.

J UNCACEAE

Juncus BuFronius L. Along the lower trails.
Juncus MAcER S. F. Gray. Baker Lake region.
Luzuxa campeEstrE (L.) DC. In open places in lower forests.

LILIACEAE

CaMASSIA QUAMASH (Pursh) Greene. Rare on grassy slopes.
DisporUM OREGANUM (Wats.) Benth. and Hook. Lower forest.
Disporum Smiruit (Hook.) Piper. Baker Lake region.
FRITILLARIA LANCEOLATA Pursh. Alpine meadows.

Luoyp1a seroTina (L.) Sweet. Moist ledges near timber line.

OrCHIDACEAE

CoRALLORHIZA MACULATA Raf. Lower forest.
HaABENARIA DILATATA Pursh. In swamp about Baker Lake.

SALICACEAE

Poputus TREMULOWES Michx. Lower slopes.

Poprutus TricHocarpa Torr. and Gray. Baker Lake and along banks of larger
streams.

Sax ARTICA Pall. var. suscorpatTa (And.) Schn. Skyline Ridge, J. W. Thomp-
son.

SALIX CANADENSIS Cock. Alpine meadows, 6000—7000 feet elevation.

SALIX LASIANDRA Benth. var. Lyatiu Sarg. Baker Lake region.

Satix nivaLtis Hook. Skyline Ridge, J. W. Thompson.

SALIX PETROPHILA Rydb. var. caEspirosA (Kenn.) Schn. Skyline Ridge, J. W.
Thompson.

Sauix ScouLer1aANA Barr. Lower slopes.

SALIX SITCHENSIS Sanson. Open forests.

BETULACEAE
BeETULA OCCIDENTALIS Hook. Common on lower slopes.

LoraANTHACEAE .
ARCEUTHOBIUM CAMPYLOPODUM var. TsucENsIs (Rosendahl) Gill. Forming
“witches brooms” associated with Tsuga heterophylla (Raf.) Sarg. and
T. mertensiana (Bong.) Sarg.

POLYGON ACEAE
PoLyGONUM AVICULARE L. A weed along the lower trails.
Potyconum Convotvutus L. <A weed above the village of Glacier.
Potyconum Dovetast Greene. Dry slopes.
Potyconum Hynprorrper L. Along ditch banks.
Potyconum Persicaria L. A weed along the lower roads.
PoLtyconvuM viviparum L. Alpine meadows above 6000 ft. elevation on Skyline
Ridge. Also reported by J. W. Thompson.
RumeEx crispus L. <A weed in the lower grassy places. |

CHENOPODIACEAE
CHENOPODIUM ALBUM L. A weed about Glacier. |

AMARANTHACEAE
AMARANTHUS RETROFLEXUS L. A weed about Glacier.

1938] MUENSCHER: FLORA OF MOUNT BAKER 267

CARYOPHYLLACEAE

ARENARIA MACROPHYLLA Hook. Open places in lower forest.
ARENARIA SERPYLLIFOLIA L. On sandy stream banks.
Crrastium viscosum L. Grassy banks, lower forest.
CERASTIUM vuULGATUM L. Grassy slopes, lower forest.
SILENE NocTiFLorA L. <A roadside weed about Glacier.
SPERGULA ARVENSIS L. A weed along lower roads.
SPERGULARIA RUBRA (L.) Presl. Baker Lake.

STELLARIA MEDIA (L.) Cyrill. A common weed about Glacier.

RANUNCULACEAE

ANEMONE Lyatiu Britt. On wooded ridges near timber line.

De.treHinium Mewnziesit DC. Grassy slopes in lower meadows.

Ranvuncvuuus acris L. Roadside weed near Glacier.

RANUNCULUS AQuUATILIS L. var. cAPILLACEUS DC. Submerged in Baker Lake.

Ranvuncutvus Boncarpi Greene. Lower forests.

Ranvuncuutus Macounnr Britt. Shore of Baker Lake.

RANUNCULUS ORTHORHYNCHUS Hook. Table Mountain, J. W. Thompson 9738.

RANUNCULUS REPENS L. A weed along roads and trails.

RANUNCULUS REPTANS L. var. ovauis (Bigel.) Torr. and Gray. Shore of Baker
Lake.

THALICTRUM OCCIDENTALE Gray. Common on moist slopes in meadows.

BERBERIDACEAE

ACHLYs TRIPHYLLA (Smith) DC. Common in lower forests.
BERBERIS AQUIFOLIA Pursh. Gravelly slopes above Glacier.

CRUCIFERAE

ARaABIs GLABRA (L.) Bernh. Near Glacier, J. W. Thompson.

BarBAREA ORTHOCEROS Var. DOLICARPA Fern. Boggy places along streams.

Brassica CAMPESTRIS L. A weed.

CapseLLa Bursa-pastoris L. A weed along lower roads.

DraBa PREALTA Greene. Skyline Ridge, J. W. Thompson.

Drasa sTENOLOBA Ledeb. High ridges and grassy ledges.

ERYSIMUM CHEIRANTHOIDES L. A weed along lower roads and trails.

Lepinium MepIuM Greene. Shore of Baker Lake.

Rorgipa Hispipa (Desv.) Britt. var. GLrasratTa Lunell. Wet banks along lower
streams and ponds.

SISYMBRIUM ALTIssIMUM L. A weed.

SISYMBRIUM OFFICINALE Scop. A weed.

SMELOWSKIA OvALIs Jones. Heliotrope Ridge, J. W. Thompson.

Supunaria aquatica L. Submerged in Baker Lake and Baker River.

CRASSULACEAE

SEDUM INTEGRIFOLIUM (Raf.) A. Nels. Moist ledges, 6000-7000 feet elevation,
Skyline Ridge, Heliotrope Ridge.

SAXIFRAGACEAE

PHILADELPHUs GorDONIANUs Lindl. Open areas in lower forest.

RIBES SANGUINEUM Pursh. Burned-over lower slopes.

SAXIFRAGA OPPOSITIFOLIA L. Skyline Ridge, J. W. Thompson.

SAXIFRAGA RIVULARIS L. Skyline Ridge, J. W. Thompson.

SUKSDORFIA RANUNCULIFOLIA (Hook.) Engler. Moist cliffs, J. W. Thompson.

TIARELLA LACINIATA Hook. Lower forests of Baker Lake region. Perhaps this
is only a form of T. trifoliata with deeply cleft leaves.

RosACEAE
Fracaria vesca L. Grassy slopes near Glacier.
Matus Fusca (Raf.) Schneid. Swamps along lower streams.
Puysocarpus OpuLiFoLius (L.) Maxim. Baker Lake.

268 MADRONO [Vol. 4

POTENTILLA GLAUCOPHYLLA Lehm. Alpine meadows.

PoTeNnTILLA NorvVEGICA L. var. HIrsuTA (Michx.) Lehm. Grassy slopes.
PoreNTILLA PaLustris L. Marsh about Baker Lake.

Prunus avium L. Escaped on burned-over slopes above Glacier.
Rosa GymnocarPa Nutt. Common in open forests.

Rosa NuTKANA Presl. Baker Lake.

Rvusus NivAuis Dougl. On slopes in lower forest.

SprraEA Dovuciastt Hook. Baker Lake region.

SpPmaAEA LUCIDA Dougl. On gravelly slopes near Glacier.

SPIRAEA ROSEATA Rydb. Springy bank, below Austin Pass.

LEGUMINOSAE

LatHyrus pALustris L. var. Myrtirotius (Muhl.) Gray. Common on burned-
over slopes.

Lupinus Lyattiu Gray. Exposed ridges about 7000 feet elevation.

Mepicaco Lurutina L. Escaped along the lower roads and trails.

TRIFOLIUM DUBIUM Smith. Escaped along the lower roads and trails.

TrRIFOLIUM HyBRIDUM L. Escaped along the lower roads and trails.

TRIFOLIUM PRATENSE L. Escaped along the lower roads and trails.

TRIFOLIUM REPENS L. Escaped along the lower roads and trails.

VictA AMERICANA Muhl. Baker Lake region.

GERANIACEAE

Eropium cicutartum (L.) L’Her. Along trails about Glacier.
GERANIUM MOLLE L. Grassy places along lower roads.

CALLITRICHACEAE
CALLITRICHE PALUSTRIS L. Baker Lake.

ACERACEAE
Acer GLABRUM Torr. var. Dovuciastt Hook. Moist slopes and edges of lower
meadows.
RHAMNACEAE

RuHAMNUs PursHIANA DC. Open lower forest.

HALORRHAGIDACEAE
Hiprvurvus vutearis L. Baker Lake; Baker River.

ONAGRACEAE

EPiILosiuM MINUTUM Lindl. Dry slopes and lower meadows.
EpiLosium PANICULATUM Nutt. Open places, Baker Lake.

UMBELLIFERAE

CICUTA OCCIDENTALIS Greene. In marshes along lower streams.
Daucus Canora L. Roadside above Glacier.

OENANTHE SARMENTOSA Pres]. Common in marshes in lower forest.
Osmorriiza optusA (Coult. and Rose) Fern. Moist banks.
Pastinaca sativa L. Along roadside, Glacier.

SANICULA SEPTENTRIONALIS Greene. Rocky slopes in meadows.

ERICACEAE

GAULTHERIA HUMIFUSA (Graham) Rydb. Rocky slope on Table Mountain.
MoneEseEs UNIFLORA (L.) Gray. Lower forest.

Monortropa UNIFLoRA L.. Lower forest. |
PyRroLA CHLORANTHA Swartz. Lower forest.

PRIMULACEAE
LysIMACHIA THYRSIFLORA L. Marshes about Baker Lake.

1938 ] MUENSCHER: FLORA OF MOUNT BAKER 269

GENTIANACEAE

GenTIANA AMARELLA L. Grassy slopes in meadows.
MENYANTHES TRIFOLIATA L. Marshes about Baker Lake.

PoLEMONIACEAE

CoLLOMIA HETEROPHYLLA Hook. Dry banks in lower forest.
CoLtomia LINEARIS Nutt. Open places in lower meadows.
Microsteris GRacitis (Dougl.) Greene. Dry slopes.

HypROPHYLLACEAE
NEMOPHILA PARVIFLORA Dougl. Along stream banks in lower forest.

LABIATAE

Gateopsis Tetranit L. Baker Lake region.

Lycorus uniFLorus Michx. Shores of Baker Lake.

Mentua arvensis L. var. caNnaDeNsIs (L.) Brig. Marshy places in lower forest.
MENTHA ARVENSIS L. var. GLABRATA (Benth.) Fern. Shores of Baker River.
NEPETA HEDERACEA (L.) Trev. Baker Lake region.

ScUTELLARIA LATERIFLORA L. In marshes along the lower streams and lakes.
StTacHys cIL1ATA Dougl. In swamps in the lower forests.

ScROPHULARIACEAE

COLLINSIA PARVIFLORA Dougl. Open places in lower forests.
DIGITALIS PURPUREA L. Naturalized along the lower roads and trails.
Mimvutuvus curratus DC. In marshes along Baker Lake.

Mimvtus moscuatus Dougl. Moist stream banks.

VersBAscuM THapsus L. A weed along roads.

VERONICA ARVENSIS L. In grassy places along stream banks.
VERONICA SCUTELLATA L. In marshes in lower forests.

OroBANCHACEAE
OrRoBANCHE UNIFLORA (L.) Britt. Dry ledges in lower meadows.

PLANTAGINACEAE
PLANTAGO LANCEOLATA L. A weed along roads and trails.

RUBIACEAE

Gatium APARINE L. In lower forests.
Gatium CiayToni Michx. Moist lower forests.
GALIUM BIFoLIUM Wats. Among boulders in the higher meadows.

CAPRIFOLIACEAE

Lonicera citiosA Pursh. Lower forests.
LoniIcerRA UTAHENSIS Wats. Rocky ledges in upper forest, to timber line.
SympnoricarPus atgus (L.) Blake. Open places on lower slopes.

CoMPOSITAE

ADENOCAULON BICOLOR Hook. Common in lower forests.

ANTENNARIA Howe Greene. Alpine meadows.

ANTENNARIA LANATA (Hook.) Greene. On stony ridges above timber line.

ANTENNARIA TOMENTELLA A. Nels. Alpine meadows.

ANTHEMIS ARVENSIS L. <A weed in open places about Baker Lake.

ArcTIuM MINUS Bernh. A roadside weed.

ARNICA LONGIFOLIA D. C. Eaton. On wet banks along Grouse Creek.

CHRYSANTHEMUM LEUCANTHEMUM L., var. PINNATIFIDUM Lec. and Lam. A weed
about Glacier.

Crepis capityaris (L.) Wallr. In grassy places in the Baker Lake region.

Ericeron acris L. var. pestis A. Gray. Skyline Ridge, J. W. Thompson.

EriceRoN ANNUUS (L.) Pers. Weed in grassy places between Glacier and
Shuksan.

270 MADRONO [Vol. 4

ERIGERON CANADENSIS L. Weed in grassy places between Glacier and Shuksan.

ERIGERON PHILADELPHICUS L. Weed in grassy places between Glacier and
Shuksan.

ERI0oPHYLLUM LANATUM (Pursh) Forbes. On dry ledges in the lower meadows.

GNAPHALIUM PURPUREUM Nutt. In gravelly soils, Baker Lake region.

GNAPHALIUM ULIGINOsuM L. Along lower trails and stream banks.

HypocHaeERis RADICATA L. <A weed in grassy places about Glacier.

LactTuca scaRioLA L. var. INTEGRATA Gren. and Godr. A weed of disturbed
places.

Lacruca spicata (Lam.) Hitche. Moist ground near Glacier, J. W. Thompson.

Manta cromerata Hook. A weed along lower trails and roads.

MatRricaRIA SUAVEOLENS (Pursh) Buch. A weed about Glacier.

Senecio Eximert Piper. Rocky ledges near perpetual ice above Heliotrope
Ridge.

SENECIO PAUCIFLORUS Pursh var. FALLAx Greenm. In open places in the lower
forests.

SENECIo vutcARIs L. A weed along the lower roads.

Sotipaco LeEpIDA DC. var. FaLLAx Fern. Open places between Glacier and
Shuksan.

Soncuus oLerAceus L. A weed about Glacier and Baker Lake.

TARAXACUM OFFICINALE Weber. A weed in grassy places along lower roads and
trails. ,

Cornell University,
Ithaca, New York, January, 1938.

LITERATURE CITED

1. Muenscuer, W.C. Flora of Whatcom County, Washington. Muhlenbergia
9: 101-132. 1914.

2. ———_—_———.. Review: St. John, Harold, and Edith Hardin. Flora of
Mount Baker. Torreya 31:15. 1931.

3. St. Joun, Harowp, and Epiru Harpin. Flora of Mount Baker. Mazama
11: 52-102. 1929.

4. THomrson, J. Wm. 1930 Washington Fern Notes. Amer. Fern Journ. 21:
117-124. 1931.

5. ————————. Notes on the Flora of the State of Washington, I. Rhodora
34: 236-239. 1932.
6. ———————_. Notes on the Flora of the State of Washington, III. Rhodora

37: 417-425. 1935.

NOTES ON STACHYS RIGIDA NUTT.

Cart EPLinG

While botanizing in Del Norte County, California, in 1935,
the author found at two stations near Gasquet an unfamiliar form
of Stachys, apparently referable to S. rigida Nutt. This has since
been collected in two other places and since it appears to be a
reasonably stable geographic race it is here described as new.
The specimens cited are deposited in the Herbarium of the Uni-
versity of California at Los Angeles.

Stacuys ricipa subsp. lanata subsp. nov. Herba substricta
altitudine 25-830 cm. superne frequenter ramosa caulibus pilis
mollibus subretrorsis dense hirsutulis; foliorum laminis oblongis
5—7 cm. longis 12-24 mm. latis utrimque praesertim subtus lanatis
mediorum petiolis 3-10 mm. longis elatis; spicis subcongestis
lanatis 5-10 cm. longis.

1938] EPLING: STACHYS RIGIDA 271

CatirorniA. Del Norte County: Bear Basin near Gasquet,
August, 1935, C. §& R. Epling, type; Patrick’s Creek, altitude 1000
feet, July 21, 19387, Parks §& Tracy 11405; from Smith River Can-
yon up to rocky hills at 2000 feet altitude, Smith River and
Patrick’s Creek, July, 1937, H. E. §& S. T. Parks 24024; French
Hill, near Gasquet, August, 1935, C. § R. Epling.

Following is a key which will serve approximately to segre-
gate the subspecies of Stachys rigida Nutt:

Petioles of lower leaves usually 2.5-4 cm. long.
Montane plants of the North Coast and
Cascade ranges (reappearing in
San Diego and Riverside coun-
ties) ; commonly 60 to 100 cm. tall,
leaf-blades tending to oblong or
deltoid-oblong .................. S. rigida subsp. typica
Plants chiefly of the coast ranges south
of San Francisco Bay and of south-
ern California, but growing as far
north as Dyerville, Humboldt
County, and as far inland as Wil-
low Creek, Humboldt County; com-
monly less than 60 cm. tall, leaf-
blades tending to ovate or cordate. S.rigida subsp. quercetorum
Petioles of the lower leaves usually less than 2.5
cm. long; leaf-blades usually narrowly ob-
long or oblong-ovate; plants chiefly of the
Sierra Nevada and Del Norte County.
Both leaf surfaces softly lanate, silvery. 8S. rigida subsp. lanata
Both leaf surfaces glabrate and green,
or at most thinly hirsute .......... S. rigida subsp. rivularis

Stachys rigida subsp. typica is found frequently in the Shasta
plateau and extends northward to the Columbia River in the Cas-
cade Mountains. It occurs in fairly typical form as far south as
Plumas and Butte counties, but merges in this region with subsp.
rivularis which is characteristic of the Sierra Nevada. The speci-
mens from the Warner Mountains of Modoc County which I have
referred to the latter subspecies are very close to S. pilosa Nutt.,
the Rocky Mountain homolog of S. palustris L. Subspecies rivu-
laris is also found in Napa and Lake counties.

Stachys rigida subsp. quercetorum is found from Lower Cali-
fornia to southern Oregon, usually at lower elevations, and as the
name implies, is often associated with the oak woodland. It is
readily recognized south of San Francisco Bay, and may be dis-
tinguished from S. bullata Benth. by the oblique and prominent
annulus. North of San Francisco Bay, however, it seemingly
passes into three other forms: S. Emersoni, S. rigida subsp. typica
and subsp. lanata. In this region it is generally a coarser plant
of larger parts than farther south and is much more tolerant of
shade, occurring in almost complete shade in the redwood groves
of Humboldt County, for example, in the Bull Creek Grove. In
Del Norte County, it is apparently replaced in the redwoods by

272 MADRONO [Vol. 4

a shade form of Stachys Emersoni. However, plants are found
frequently in Humboldt and Mendocino counties which are diffi-
cult to assign certainly to either of these species. In Humboldt
County subsp. quercetorum is transitional to subsp. typica and in
Mendocino and Sonoma counties forms occur which in turn sug-
gest the silvery dense pubescence of subsp. lanata, but have the
habit of subsp. quercetorum.

Stachys Emersoni is by no means a homogeneous species, and
indeed, appears to partake of characteristics both of S. bullata
and S. ciliata. ‘The flowers are characteristically quite dark rose
purple. As the author has indicated elsewhere there is reason to
believe that S. Riederi of Chamisso described from “‘Kamtschatka”’
is conspecific with this species. The type of S. Riederi has not
been located. Recent examination of the Labiatae of the Mocifio
and Sessé herbarium has further shown that S. meaicana of
Bentham, a “‘lost’’ species, is certainly conspecific with S. Emer-
soni. The specimen of Mocifio and Sessé is very similar to that
collected by Abrams (no. 11246) at Ilwaco, Washington.

University of California
at Los Angeles, April, 1938.

EREMOCARPUS BENTHAM: PREOCCUPIED?

Lovis C. WHEELER

The name Eremocarpus was proposed by Bentham (Bot. Voy.
Sulphur 53, pl. 26, 1844) for a monotypic genus of Euphorbiaceae.
The validity of the generic name was questioned by Coville
(Contr. U. S. Nat. Herb. 4: 194. 1893) on the ground that
“Eremocarpus was first used by Reichenbach, in 1837, as a desig-
nation for a genus of Hypericaceae.”’ Piper, apparently accept-
ing Coville’s statement on faith (Contr. U. S. Nat. Herb. 11: 382.
1906), renamed Eremocarpus Bentham as Piscaria. The alleged
Eremocarpus of Reichenbach which first appeared in his synopsis
of the Hypericaceae (Handbuch Nat. Pflanzensystems 307. 1837)
is there credited to Spach. The context makes it evident that
Reichenbach merely suffered a lapsus memoriae regarding Eremo-
sporus Spach (Hist. Nat. Veg. 5: 342. 1836, nomen nudum; Ann.
Sci. Nat. Bot. ser. 2, 5: 855, 1836, Conspectus Monogr. Hyperi-
cacearum 349-369). Reichenbach published Eremocarpus as
follows:

“q) Drosautheae: [sic, error for Drosantheae] capsula tri-
cocca, coccis 1—3-spermis, demum cum placenta deciduis. Eremo-
carpus Spach. Drosanthe Spach.”

It is quite evident from the similarity of the names and de-
scriptions and from the mention of Spach’s monograph (Hand-
buch Nat. Pflanzensystems 308) that Reichenbach based his char-
acterization of Drosantheae on the diagnosis of “Sectio I. Dro-

1938] BRACELIN: YNES MEXIA 273

santhineae” Spach (Ann. Sci. Nat. Bot. ser. 2, 5: 354) under
which Spach included only Eremosporus and Drosanthe. Without
description and without indication of any intent to rename,
Reichenbach in a later publication again mentions “Eremocarpus
Spach” (Rep. Herb. Nomenclator 210. 1841). Thus it seems
clear that Reichenbach was following Spach’s work and merely
changed Eremosporus Spach accidentally. The reasonable course
to follow in this case is to consider “EHremocarpus Spach” as an
unintentional change without any power to preoccupy. The
International Rules of Botanical Nomenclature do not provide for
such dubious cases. If anyone wishes to claim that this was an
intentional renaming and, though illegitimate, capable of pre-
occupying the name, it is only necessary to enforce strictly the
requirement for valid publication in Article 42: “(2) by the cita-
tion of a previously and effectively published description of the
genus under another name’’; this will automatically exclude
“Eremocarpus Spach.” In no case did Reichenbach give any page
reference which is certainly an essential part of a citation.
Reichenbach’s vague mention on a succeeding page (vide supra)
of Spach’s monograph is not a citation.

There is an exactly parallel case which is apropos here.
“Eremocarpus Bunge” appears in a bare list of genera (Lindley,
Veg. Kingdom ed. 2: 778. 1847). The authors of Index Kew-
ensis are probably quite right in interpreting this as Eremo-
daucus Bunge, from which it must have arisen by a lapsus.

Conclusion: Eremocarpus Bentham is not preoccupied.

Gray Herbarium, Harvard University,
Cambridge, Massachusetts,
April 5, 1938.

YNES MEXIA

Ynes Mexia, the daughter of General Enrique A. and Sarah
R. (Wilmer) Mexia, was born May 24, 1870, in Georgetown,
Washington, D.C. Her father, the son of José Antonio Mexia (a
Mexican general under President Santa Anna) was at that time
resident in Washington as a representative of the Mexican gov-
ernment. Her mother, Sarah R. Wilmer of Maryland, was of the
family of Samuel Eccleston, Fifth Archbishop of Baltimore. A
large part of her childhood was spent in Texas where the family
owned an eleven league grant upon which the town of Mexia,
Limestone County, is now located. Her early education was ob-
tained mainly in private schools in Philadelphia and Ontario,
Canada. Later, she attended St. Joseph’s College, Emmetsburg,
Maryland, and the University of California, Berkeley. She was
married in Mexico to Agustin A. de Reygadas but later resumed
the use of her maiden name. For considerable periods during
the earlier part of her life she lived in Mexico but for the past
thirty years has been a resident of San Francisco.

274 MADRONO [Vol. 4

Mrs. Mexia’s interest in botanical collecting began in 1922
when she joined an expedition led by Mr. E. L. Furlong, then
Curator of Paleontology, University of California, Berkeley; on
this occasion, however, she
made only a few permanent col-
lections. Her first important
collecting was done on a second
expedition to Mexico in 1925 in
company with Mrs. Roxana S.
Ferris, of Dudley Herbarium,
Stanford University. During
the thirteen years following she
made three additional expedi-
tions to Mexico, one to Alaska,
and two to South America, col-
lecting a total of about 8800
numbers, approximating 145,-
000 specimens. She was col-
lecting in the mountains of the
State of Oaxaca, Mexico, in
1938 when she became ill and
was obliged to return home.
Her health did not improve and
death followed on July 12, 1938.

Mrs. Mexia’s collections were
always carefully prepared and
her field notes unusually detailed. Many of the regions she
visited had been but little explored botanically and although
studies upon her collections are not yet completed they have
yielded a large number of species new to science. At present
there have been described two new genera, Mezianthus meaxicanus
Robinson (Compositae) and Spumula quadrifida Mains (Pucci-
niaceae) and about 500 new species (mostly spermatophytes) of
which more than 50 have been named in her honor. A brief
résumé of her collecting expeditions with approximate numbers
of specimens obtained is given at the end of this article.

Published accounts of Mrs. Mexia’s expeditions and special
reports upon her collections have appeared as follows: Ynes
Mexia, Botanical Trails in Old Mexico—the Lure of the Un-
known (Maprono 1: 227-238. 1929); Three Thousand Miles up
the Amazon (Sierra Club Bulletin, 1983); Camping near the
Equator (Sierra Club Bulletin, 1937); Edwin B. Bartram, Mosses
of Western Mexico Collected by Mrs. Ynes Mexia (Jour. Wash.
Acad. Sci. 18: 577-582. 1928); E. B. Copeland, Brazilian Ferns
Collected by Ynes Mexia (Univ. Calif. Publ. Bot. 17: 23-50, pls.
1-8. 1932); Mrs. H. P. Bracelin, Itinerary of Ynes Mexia in
South America (MaproNo 8: 174-176. 1935).

Fig. 1. Ynes Mexia.

1938] BRACELIN: YNES MEXIA 275

In the San Francisco Bay region Mrs. Mexia was well known
as a lecturer, having appeared before many scientific organiza-
tions. Her accounts of botanical explorations were vivid and
entertaining, and because of her skill in photography, were un-
usually well illustrated with views of the general topography and
plant associations of the regions visited.

Mrs. Mexia has been a member of the California Botanical
Society since 1915. She was a member also of the Sierra Club,
the Audubon Association of the Pacific, the Sociedad Geographica
de Lima, Peru, a life member of the California Academy of Sci-
ences, and an honorary member of Departamento Forestal y de
Caza y Pesco of Mexico.—Mrs. H. P. Bracetin, Berkeley, Cali-

fornia.

BorTranicAL EXPEDITIONS OF YNES MEXIA
Mexico

Western Mexico: September 15 to November 19, 1925; Sinaloa; expedition
with Roxana S. Ferris, Dudley Herbarium, Stanford University; 500 numbers,
3500 specimens.

Western Mexico: September, 1926 to April, 1927; states of Sinaloa, Nayarit,
Jalisco to 6000 feet elevation in Sierra Madre; 1600 numbers, 33,000 specimens.

Northern and central Mexico: May to July inclusive, 1929; Chihuahua,
Mexico, Puebla, Hidalgo; expedition led by Mr. E. L. Furlong, Department of
Paleontology, California Institute of Technology, Pasadena; 315 numbers, 5000
specimens.

Southwestern Mexico: October 31, 1937 to May 20, 1938; states of Oaxaca
and Guerrero; 700 numbers, 13,000 specimens.

ALASKA

Mt. McKinley National Park: June to September, 1938; first general collec-
tion of the Park flora; 365 numbers, 6100 specimens.

SoutH AMERICA

Brazil and Peru: November, 1929 to March, 1932, inclusive; Brazil, Rio de
Janeiro, Vicosa and Diamantina, state of Minas Geraes; Amazon and other
river courses in states of Parad and Amazonas; Transandean Peru, upper Amazon
and Santiago river valleys, Departamento de Loreto; accompanied for a short
time by Agnes M. Chase, Division of Agrostology, United States Department of
Agriculture; 3200 numbers, 65,000 specimens.

Ecuador: September, 1934 to September, 1935; coastal plains and eastern
Amazonian slope of Andes, northern highlands and Columbian border; expedi-
tion for the Bureau of Plant Introduction and Exploration, United States De-
partment of Agriculture to search for palms, cinchonas and soil-binding plants
and to make a general collection; 900 numbers, 5000 specimens. —

Peru, Bolivia, north central Argentina and Chile: October, 1935 to January,
1936, inclusive; Andean highlands; expedition of the University of California
Botanical Garden led by Dr. T. H. Goodspeed; 300 numbers, 1900 specimens.

Peru, Chile, Argentina and Ecuador: January, 1936 to January, 1937, inclu-
sive; southern Chile, Straits of Magellan, Tierra del Fuego; Peru, Cuzco,
Machu Pichu, Cerro del Pasco; Ecuador, Esmeraldas; 1000 numbers, 13,000
specimens.

276 MADRONO [Vol. 4

VIAE FELICITATIS: THE BEGINNING YEARS OF THE
CALIFORNIA BOTANICAL SOCIETY?

Wiis Linn JEPSON

Mr. Toastmaster and old-time friend: Doctor Foxworthy, Past
President of the Society, I salute you: Mr. President of the Soci-
ety: Members and Guests.

At this silver jubilee happiness for good reason pervades this
large gathering. It is a sign of continued vitality that the Cali-
fornia Botanical Society this year elected as its president, Pro-
fessor Howard E. McMinn of Mills College. It is also a source
of gratification to me personally to find him occupying the presi-
dent’s chair. Naturally my mind, tonight, goes back to the very
beginnings of the Society in 1913, and various happy reflections
arise in memory. The backbone of the Society, the most eligible
and vital part of the membership, in the earliest years, consisted
in the main of those who were not professional botanists in the
strictest sense, but men in other fields, most often in fields of
applied botany such as agriculture, horticulture, agronomy, silvi-
culture or forestry; but also including men in medicine, dentistry
and pharmacy and various business occupations. There was
reason for this. It was partly because there were extremely few
professional botanists in California at that time, but there was
another reason to which I shall advert later. Practically all of
these persons who gave the Society its most solid support, and
many others equally valuable as members, had a college bache-
lor’s degree in botany, or a master’s degree, or a doctor’s degree.
It will be interesting to recall a few that come to mind. Profes-
sor William T. Horne, now of the Citrus Experiment Station, a
plant pathologist with the soundest kind of training; Professor
W. W. Mackie, of the College of Agriculture, an able agronomist
and world traveler; Dr. H. J. Webber, of the College of Agricul-
ture, horticulturist and geneticist, a man with a distinguished sci-
entific record who was the discoverer of motile spermatozoids in

1 Response at the Annual Dinner of the Society held in Berkeley on April
23, 1938. The Toastmaster was Dr. W. W. Robbins, College of Agriculture at
Davis. Past President F. W. Foxworthy, long a forest officer in the Federated
Malay States, is now a resident of Berkeley. The President of the Society,
Professor H. E. McMinn, introduced the Toastmaster. Those botanists from
a distance included Dr. C. B. Wolf, Botanist at the Rancho Santa Ana Botanic
Garden, Anaheim; Mr. Elmer I. Applegate, Palo Alto; Dr. Carl Epling, Asso-
ciate Professor of Botany in the University of California at Los Angeles; Mr.
M. Van Rensselaer, Director of the Blaksley Botanic Garden at Santa Bar-
bara; Dr. H. F. Copeland, Professor of Botany in the Sacramento Junior Col-
lege; Dr. L. H. Knoche of San Jose; Professor D. H. Campbell, Professor Geo.
J. Peirce, Professor L. R. Abrams, Professor Ira L. Wiggins and Professor
L. R. Blinks of Stanford University; Dr. E. B. Copeland of Chico; Mr. H. A.
Dutton of Los Altos; and Dr. D. D. Keck of the Carnegie Institution at Stan-
ford. The major portion of the members present at the dinner came from San
Francisco, Burlingame, Oakland and Berkeley. Professor Peirce, a Past Presi-
dent of the Society, read the congratulatory messages from absent members.

1938] JEPSON: CALIFORNIA BOTANICAL SOCIETY 277

Zamia, a genus of cycads; Dr. Herbert M. Evans, of the School of
Medicine of the University of California, who never forsook his
early love for botany; Elizabeth Smith, plant pathologist in the
College of Agriculture, one-time officer of the Society; Professor
P. B. Kennedy, a skillful agronomist, who had his doctor’s degree
in botany from Cornell University, and who was for a time presi-
dent of the Society; Albert Walker, a pharmacist, who always
increased the value of field trips by his knowledge of applied
botany as did Dr. A. W. Card, a dentist; F. W. Koch, who was
-then in charge of the department of biology in the Galileo High
School at San Francisco, added his store of field knowledge to the
Society’s interests; in his youth he had been attached in a minor
capacity to the Death Valley Expedition of the United States
Biological Survey and never forgot his field experiences with C.
Hart Merriam and F. V. Coville; Frederic T. Bioletti, viticulturist
in the College of Agriculture, who always remembered vividly
the botanical collecting trips of his early days and thrilling dis-
coveries of new species. One of the most valuable and loyal
members was Dr. W. C. Blasdale. Although his profession was
that of a professor of chemistry in the University of California,
botany was with him a capital hobby; he had done much field
work and his published papers on botanical subjects are critical
contributions to their field. Many, many others could be men-
tioned if there were time to list them.

The first secretary of the Society was Mrs. D. W. de Veer.
If wisdom be the art of so using knowledge or facts as to produce
the most satisfactory results, Mrs. de Veer had much wisdom and
handled our affairs to advantage. The numerous field trips
scheduled by the Society in that early day necessitated a great
deal of clerical work. On account of our extremely limited
funds it was a rigid rule that all this clerical work must be done
by members. Sometimes volunteers were ready, at other times
perhaps not so ready. Sometimes I would make appeal in the
name of the great Swedish botanist, Linnaeus, or of Theophrastus
the Greek, without finding my office door blockaded by helpers.

The first person in these United States to put the unemployed
to work was Tom Sawyer. The original Tom Sawyer of the Cali-
fornia Botanical Society was our second secretary, Miss Anna
Ehlers, who had been a student under that great teacher of bot-
any, Professor Charles E. Bessey of the University of Nebraska.
She had not the slightest difficulty in putting members to work.
Indeed, it was a byword of that time that she made them work
like Trojans. During my long tenure as president of the Society,
the secretary of the Society in her capacity as secretary would
often look at me reproachfully on account of my numerous short-
comings and perhaps I had reason, therefore, to be thankful that
Miss Ehler’s disposition was entirely amiable. The next secre-
tary was Professor H. E. McMinn of Mills College, but inasmuch
as he lived some distance away from our headquarters it was Mrs.

278 MADRONO [Vol. 4

Adeline Frederick, Assistant Secretary of the Society, who did
most of the secretarial work. She brought to the task a remark-
able sense of perception and a sagacity and soundness of judg-
ment that furthered the Society's progress in stabilizing ways
that are observable to this day. Succeeding Mrs. Frederick as
secretary was Mrs. Linda Dodd. It was her function, on account
of the years in which she served, to pour oil on troubled waters.
She successfully smoothed out many a rough spot and saved us
many an inconvenience. .

Another early member, for a time an officer of the Society,
was Dr. C. B. Bradley, Professor of Rhetoric. We spoke of him
as a good man gone wrong. In his early years he had a few
happy weeks in the herbarium of Asa Gray and missed becoming
a botanist by only a narrow margin. Never, though choosing the
field of English, did he lose his early interest in botanical science.
His published papers on botanical subjects are soundly, carefully
and conscientiously done, and are true scientific contributions.
In questions of taste and language he was our undisputed arbiter,
especially in problems of transliteration of Greek roots into Latin
or directly into English. If there were doubt about a Greek
diphthong, as to whether transliteration should be literal or other-
wise, we accepted his judgment as infallible. There is an anec-
dote concerning him, involving a curious case of transliteration,
which I would tell if ladies were not present. Oh, well, Ill tell
it anyway. Women should not become members of the California
Botanical Society unless they are willing to take the risks of the
Society. One summer Professor Bradley was in the Sierra Ne-
vada with a small party of young men, naturalists all. One day,
being unwell, he elected to stay in camp and write a letter. The
cook of the party was a foreigner of the lower class whose
English vocabulary was extremely limited but who thought he
understood what was said to him. When the cook reported to
Professor Bradley for orders, in that careful and precise speech
for which he was famous, Professor Bradley said to him: “A cup
of tea is all I wish; that will fully meet my requirements. As for
the young gentlemen they must speak for themselves. I could
not order for them.” The cook was a little bewildered by this,
but thought he understood fully. When the ravenously hungry
field crew came into camp and found no food ready, the cook
was taken to task but he defended himself. “But,” said one,
“didn’t Professor Bradley tell you to get up our grub?” “No,”
said the cook. ‘But he told you something; what did he say?”
Said the cook: “He said, ‘By Jinx, I’m all right, but them
blankety blankety guys can go blankety blank.’ ’’ When Professor
Bradley was told of this case of transliteration, he enjoyed it as-
hugely as anyone else, for he had a deal within him of the live-
liest fun and humor.

In the year 1901 there called upon me at my laboratory in
Berkeley a research botanist who had come to the Pacific Coast

1938] JEPSON: CALIFORNIA BOTANICAL SOCIETY 279

to investigate botanically the sources of the drug cascara sagrada,
which is derived from the tree, Rhamnus Purshiana. He had been
sent by the Wellcombe Research Laboratories of the great Lon-
don drug and chemical firm, Burroughs and Wellcombe. This
botanist’s name was P. E. F. Perrédés. In his surveys up and
down the Pacific Coast he accumulated a store of highly inter-
esting facts regarding our species of Rhamnus, the amount of vari-
ation in Rhamnus Purshiana as it ranged south into California, the
places where it interbred with Rhamnus californica, and the extent
to which the bark supply was adulterated by Rhamnus californica
bark, which had quite the same cathartic properties but different
therapeutic values. I extended to Perrédés every hospitality and
aid, and he was most generous in sharing his knowledge; but it
struck me that in such a case as this the hospitality of any one
individual was inadequate and that there should be a botanical
society in order to fully honor him. Perrédés was a jolly soul
and vastly interested in the social habits and ways of my country-
men. One day he came into my laboratory and exclaimed very
genially: “You Americans are a very curious people!” I replied,
“Certainly we are. Go ahead.” ‘You are a very strange people,”
he went on. “I observe that you sweep your parlors with a
lawn-mower; you take up the collect in church with a fry-pan;
you brush your clothes with a broom; and bless my soul, you call
a pram a baby-carriage.” There were before and after this many
other foreign botanists, some of them highly distinguished men,
who visited California in those years, botanists from Europe,
Australia, New Zealand, Japan and South Africa. In my very
early years at Berkeley one of these visitors was Dr. Paul Knuth
from Kiel, Germany, the author at a later time of a monumental
“Handbook of Flowers and Pollination.” Of him I still have a
lively picture in my memory, as dressed in a tropical suit he ran
hither and thither in the Garden of Native Plants capturing in-
sects with his net. Such travelers continued to appear in Cali-
fornia and it was constantly borne in upon me that there should
be a botanical society to greet them, honor them and benefit by
their presence.

As planned it was determined in my mind that such a botani-
cal society must needs envision four things: 1. It must be a state-
wide botanical society. 2. Its field of research must be the whole
of western America. 38. It must have a printed journal in order
to keep touch with its scattered membership. 4. It must have
world relations and a world point of view and keep abreast of the
progress of botanical science in every land by honoring in its
public meetings botanical travelers from all parts of the earth
who would interpret for us the research results of their country.
These were the four primary objectives.

By means that were highly fortunate the Society to some de-
gree realized these objectives within a few years. While the
strength of the membership was in the cities of the San Francisco

280 MADRONO [Vol. 4

Bay region, almost immediately botanists in various and often
distant parts of California joined the Society and kept up their
membership for decades, even though unable to attend a single
meeting of the Society. If these members in remote or lonely
places botanically could have a durable trace of light connecting
them with the Society by receiving the Society’s meeting notices
they were happy to lend their continued support to a botanical
organization and cooperate in its work. Mrs. Clara A. Hunt of
Saint Helena kept up her membership faithfully for years, though
never attending a Society meeting. But the inspiration derived
from membership caused her to stir up Napa Valley botanically
each year with an exhibition of native plants in the beautiful
grounds of her Saint Helena home. Mr. Milo N. Wood, pomolo-
gist of the United States Department of Agriculture, who was
stationed at Sacramento, an expert on Juglans regia, maintained
his connection with the Society indefinitely; and Mrs. Harriet P.
Kelley, at Selma in Fresno County, started a branch of the Soci-
ety at that place which flourished as long as she was resident
there.

For its journal the name Maprono was chosen, because Ma-
DRONO is euphonious, because it makes a brief and handy title,
because it is a name associated with the region, because, most of
all, it is the name of the native arbute tree which has great bio-
logical and forestal significance. The native vegetation of Cali-
fornia is not merely rich and varied to a remarkable degree; it
has also an extraordinary geographic significance and an almost
ponderous relation to geological history. In such a land a bo-
tanical journal must in every way be original and filled with
original articles. It is unthinkable that with such a physical
background, with such a geographic region as our field, that the
Society should in any way copy the journals or adopt the mores
of older lands; it is unthinkable that it should so lack in origi-
nality as to call its journal, for example, a bulletin, which word
used a thousand times means perhaps nothing, perhaps anything.
There is a great deal in a name if it be original. The name
MaproNno for our journal was an original thought happily seized
by the imagination. It means so vastly more than bulletin that
it at once puts the journal on a different plane. In addition to
the original articles a portrait of a distinguished botanist or bo-
tanical explorer having to do with botany in California was
planned as a frontispiece for each volume, thus placing upon each
volume a fitting and distinctive seal. Volume one carries as
frontispiece the portrait of Archibald Menzies, the botanist of the
Vancouver Expedition, the first botanist to collect in any marked
degree in California and whose name is inseparably associated
with many of our most characteristic plants as discoverer, and
especially with Arbutus Menziesii, the madrofio. The second vol-
ume uses as frontispiece the portrait of the distinguished botanist,
Thomas Nuttall, who came to California in 1886. It was believed

1938] JEPSON: CALIFORNIA BOTANICAL SOCIETY 281

that if the journal fortunately continued and attained to a dozen
or twenty volumes or more, this one feature alone would give it
an original interest to all botanists in every land.

The fourth condition that the society establish world relations
was auspiciously begun when five months after its first meeting
it entertained at dinner the members of the International Phyto-
geographic Excursion, which included amongst them some of the
more distinguished botanists of the earth. The party included
Dr. Adolf Engler of Berlin; Professor C. von Tubeuf of Munich;
Dr. Edward Ribel, Dr. H. Brockmann-Jerosch and Professor Carl
Schroter of Ziirich; Professor A. G. Tansley of Cambridge, En-
gland; Dr. T. J. Stomps of Amsterdam; Dr. Ove Paulsen of
Copenhagen; Dr. Carl Skottsberg then of Uppsala; Professor
H. C. Cowles of Chicago; Dr. A. Dachnowski of Columbus; Pro-
fessor and Mrs. F. E. Clements of Minneapolis; Mr. Geo. D.
Fuller of Chicago; and Dr. Geo. E. Nichols of New Haven. Over
one hundred members of the Society and their friends joined in
a demonstration of hospitality to these guests. The arrange-
ments and service were all that could be wished, the dinner being
held in a quite noble private dining room of the Hotel Oakland.

The main festival of the Society in its year of activity is the
annual dinner, which has been held regularly each year since the
first one in 1913. At this dinner it has often been possible to
have as chief speaker a visiting European or eastern botanist or
botanical traveler from some part of the earth. Greetings are
read from absent members and usually some five or six members
talk a few minutes before the main speaker comes on.

The importance of this function, extending hospitality to bota-
nists from other lands, cannot well be overestimated. The flora
of western America nor that of California is not a vegetation
apart, but is involved in its history with all other vegetation areas
of the earth. Political boundaries, of course, have no signifi-
cance. The same thought applies to botanists themselves. When
one is reading Schimpfer’s ‘“Pflanzen-geographie,’ Linnaeus’
“Fundamenta Botanica” or the “Enquiry into Plants” by Theo-
phrastus, one never thinks of these botanists as belonging to any
particular race or state but only as belonging to the company of
scientists who have in every age carried forward the light of
rationalism. Scientists in the world are a very small group, but
they are the only men in any land who have with each other a
common understanding, who are able to arrive at common judg-
ments, because they are the only men whose decisions are arrived
at purely on the basis of scientific facts, whose decisions are
purely objective. In their mental processes, motives of race,
country, creed, political orders or social emotion are wholly lack-
ing. It is, therefore, fitting that in our land of California there
should be a botanical group which does its share to tend the flame
of pure rationalism as a signal and a sign to similar groups in
other lands about the earth.

282 MADRONO [Vol. 4

Since the idea of a Society was definitely conceived in the
year 1902, some one may note the lapse between that date and
the year 1913 when the Society was founded. The answer is not
easy to express. Of professional botanists in California at that
early day there were but few outside two small groups, one at
Stanford University and the other at the University of California.
There was no intercourse between the men of these two groups.
The men at Berkeley had never called on the men at Stanford,
the men at Stanford had never come to Berkeley. The only ex-
ception to this rule was myself. I had been down to the Stan-
ford Department of Botany a few times. Eventually I decided
to go ahead with the formation of a society, believing that when
the botanists of the state generally were members these two
groups would follow. It was certainly not to be expected that
the professors at Stanford would travel the long distance to
Berkeley to join a local society, a society having to do merely
with Berkeley. It was obvious the Society must first establish
proper claims to support. That judgment as to the Stanford
group proved correct. As soon as it was possible I went down
to Stanford more frequently. It was not difficult to win to mem-
bership so good-natured and genial a man as Dr. LeRoy Abrams
and in time he was elected a vice-president of the Society. In
1918 the dean of the summer session at the University of Cali-
fornia was Walter Morris Hart, a distinguished scholar and
gentleman. I was given an opportunity to nominate the profes-
sor of botany in the summer session and I nominated Professor
George James Peirce of Stanford University. Professor Peirce
was called. He came to Berkeley for the summer, brought Mrs.
Peirce and his three charming daughters, and all the family en-
joyed themselves hugely. He came again the next year and
possibly a third year, though I have forgotten. In any event it
led to relations which were to prove of great importance to the
California Botanical Society later on. A few years later Profes-
sor Douglas Houghton Campbell became a member. This acces-
sion led to certain associations which might otherwise have been
missed.

Only yesterday I had a letter stamped by the military censor
from my niece who is in Spain. In the city where she lives the
alarm sounded, warning of the approach of a bombing plane with
its bombs dealing a terrible death to people and ghastly destruc-
tion to buildings. Immediately, with her servants she fled to the
bomb-proof shelter where she was joined by the members of a
neighboring family. The mother of this family, however, on
getting half way to the shelter remembered that a rug sent home
from the cleaners had been left rolled up on the floor. She
could not bear the thought that some one might come and find the
room untidy, and so went back, unrolled the rug properly and
then joined her family in the shelter. This is what might be
called devotion to, enthusiasm for an ideal.

1938] JEPSON: CALIFORNIA BOTANICAL SOCIETY 283

The enthusiasm of the early members of the Society for the
welfare of the Society was like that. From the very beginning
there was this deep enthusiasm. Those who joined in the Soci-
ety’s work took the greatest possible interest in it; they felt
honored to be a part of it. I recall, almost at random, the loyalty
of Inez Ray Smith of Hillsborough, the wife of Mr. James Ber-
nard Smith. She made the flora of San Mateo County her spe-
cialty and usually led the field trips scheduled for that region.
She took pains to go over the ground in advance so that the ex-
cursion might be in every way most profitable to the members
and nothing missed. I observed that in addition to her scientific
interest she had an appreciation of the beauties of nature in its
manifold forms. It is probable that this esthetic appreciation
was an inherited characteristic, since she was a cousin of the dis-
tinguished American artist, Charles Dana Gibson. Both Mr. and
Mrs. Smith enjoyed greatly their connection with the Society
which led directly to their financial aid to the Department of
Botany at Mills College and their support of the research work
of various California botanists.

As time passed the Society gradually drew into its fold bota-
nists far and near about the state and took pride in its acces-
sioned members, of whom I can only mention a few. Dr. L. H.
Knoche, who had his undergraduate training under Professor
W. R. Dudley, went to the University of Montpellier in France
to study plant geography under Professor Charles Flahault.
While in France he became interested in the problems of the
Mediterranean flora and produced his classical four-volume work,
“Flora Balearica,’ a phytogeographical study of the Balearic
Islands. Later he returned to the United States, settled once
more in his boyhood home of San Jose, built an herbarium build-
ing for his herbarium of over 100,000 specimens and a botanical
library of approximately 12,000 volumes and 35,000 pamphlets.
Milo S. Baker, another member, in his early years a well-known
explorer of northeastern California, later resided in Sonoma
County, and as professor of botany in the Santa Rosa Junior Col-
lege, brought together an admirable local herbarium and special-
ized on the genus Viola. Carlotta Case Hall, whose interests
centered on Pteridophyta and especially the ferns of California,
was another delightful member of early days. Others of note
were Dr. E. P. Meinecke, pathologist in the United States Forest
Service, and Dr. Charles Piper Smith, of the San Jose High
School, who left the eastern United States and settled in Cali-
fornia for the field advantage of his critical studies on the genus
Lupinus.

The field trips, some twenty to forty a year, were always a
very important part of the Society’s programmes in early years
and were well attended. For at least a decade, a “camping trip,”
covering two or three days in the field, was arranged for the end

284 MADRONO [Vol. 4

of May. One such excursion in 1921 was made to the Mayacamas
Range in Mendocino County where, on a high mountain side, are
situated the native plant gardens of Carl Purdy. Some one hun-
dred members formed the party, including a number of zoolo-
gists. The camp spot was situated in a most delightful vale
where the natural conditions had never been disturbed.

Such field trips were of great profit to nurserymen, horticul-
turists and foresters and to teachers in the schools, especially to
those who had recently come to California from the eastern
United States and thus found an unfamiliar flora. The values of
field experience were various and are not easy to summarize in a
few words. It sometimes happened that members who had no
particular bent for the critical side of either plant ecology or
plant taxonomy yet developed certain natural history interests of
importance. An illustrative example is that of Mrs. Ynes Mexia
whose inspirations were derived from the Society. She made
successful collecting trips to Mexico, and later, from 1929 to
1932, went to South America. On one of her expeditions she
proceeded from the eastern coast of Brazil to the headwaters of
the Amazon River and thence over the Andes, most of the way
with only Indian canoemen, to Peru. The plant materials which
she gathered on this trip are of real importance to botanists and
will permanently associate her name with certain phases of South
American botany. In addition, botanists from foreign countries
appreciated the Society’s field excursions. One recalls Dr. Alice
Scouvart, a student of Professor Jean Massart of the University
of Brussels, who, coming to live in California for a few years,
taught some of our members the values lying in the scientific
names of plants.

The field trips were unquestionably a great tonic and it were
unfortunate if they lapsed entirely. One field trip a year, planned
with elaborate care, to some remarkable area in California, would
attract attendance from the eastern United States and give us the
advantage of more frequent association with distant botanists.
There is no botanist or botanical traveler or layman interested in
the native plants but would wish to see first of all six notable
plant associations in California, namely, the Torrey pines associ-
ation on Santa Rosa Island; the Joshua tree or yucca association
on the Coolgardie Yucca Mesa in the east central Mohave Desert
north of the Calico Mountains; the big tree association in the
Giant Forest at Round Meadow above Kaweah; the serpentine
association in eastern Lake County; the redwood association of
Prairie Creek in Humboldt County; and the Bishop pine-beach
pine association on the Mendocino Coastal Plain. It were im-
possible to say which of these associations is the more remark-
able, since they are all so utterly different and are, indeed, not
comparable; but none of them is as yet botanically well known.
All await intensive study.

1938 ] JEPSON: CALIFORNIA BOTANICAL SOCIETY 285

On the Mendocino coastal plain are almost incredible sights:
bishop pine and beach pine growing to their full stature 50 to 75
feet high in loam soil; a few yards away the same species dwarfed
down to mere canes but coming into reproductive maturity and
bearing cones when only 1 to 2 or 8 feet high. A short time ago
I discovered on broken and difficult terrain a cluster of the Men-
docino cypress, a small colony of about twenty-five trees, that
probably represent the largest individuals of true cypress in the
New World. Standing before these trees one feels as if he were
in the presence of Sequoia gigantea. The trunks go on up and up
and up without sensible diminution in size to a small rounded
crown of short branches. One-eighth mile away on the gypsum
soil this same species occurs as tiny dwarfs which produce cones
when only 1 to 2 feet high.

Nor, in the planning of any major excursions, should the
great San Joaquin, where much of the primitive vegetation still
remains, be forgotten. Of all the areas surrounding the Great
Valley of California the ranges on the west side of the San Joa-
quin Valley, far south, are in summer the most desolate—as if no
living thing could ever flourish in that hopeless dryness. But
then comes the providence of winter rains in a good year and one
stands on the plain in front of the San Carlos foothills and be-
holds waterfalls of flowers down their steep faces and diadems of
yellow on the summits which run out into filagrees of gold on the
slopes and ridges. For fifteen miles north along the range, in
brilliant sunshine, one surveys that color, for fifteen miles south
the eye follows that broad band of pageantry—thirty miles of
glory. In such a year uncommon species become common and
help outline more definitely the plant associations. In such a
year, too, one may get glimpses of the relation of the plant for-
mations to the geological fault blocks. One traces a narrow
endemic to a fault line where it stops. In one case a species was
found in abundance, which had not been seen before in forty
years. It was a so-called “lost species.” The year that I was
born in California was not so far from the year in which was
published the “Origin of Species” and, as I grew up, everything
was most naturally and easily explained by various interpreta-
tions of Darwin’s beautiful theory. Some of the new species dis-
covered and published at an early day in California had never
been found again and it was explained to us that in the plentitude
of evolutionary differentiation of new species more species were
produced than could survive. Hence the “lost species” in Cali-
fornia. We have quite different ideas now. We imagine we
know more about behavior in the soil of seeds of certain native
species—dormant five, twenty, fifty or more years, until a year
comes when there is a favorable run of low temperatures in con-
nection with appropriate moisture conditions and other as yet
unappreciated or not fully understood factors.

286 MADRONO [Vol. 4

Of the early years of the Society I tend to have only happy
memories. Nevertheless, it was not always one bright and
charming holiday without untoward incident. There were days
and even years when discouragements nearly mastered the situ-
ation and I was again and again almost ready to give up. But
these things were in the end offset by progress and success and
especially by the moral support of one’s friends, by friends who
knew nothing of one’s difficulties, but whose faith was unshake-
able. Such friends were Anson and Anita Blake, Cornelius Beach
Bradley, Samuel B. Parish, Eliza B. Parish, Cornelia C. Pringle,
Frederick Hein, Emanuel Fritz and others, too many for all to be
listed now.

On one of the Society’s excursions a woman member chanced
delightedly upon a California black oak seedling in the forest and
claimed it for her garden. But she could not lift it out of the
soil, nor could two men members who came to her aid, though it
was only four inches high. They did not know that the little oak
was eight or ten years old and had been insuring its life by giving
nearly all to its root system. The California Botanical Society
was like that in its first eighteen years. It had put its all in
foundational roots. When late in 1929 the nominating committee
brought to me the list of nominees of officers for the next year I
took the list and drew a pencil through the name of Willis Linn
Jepson for president and wrote in the name of George James
Peirce of Stanford. I was now certain that the Society could not
readily be uprooted. It was ready to flourish above ground and
Professor Peirce was the one best fitted to guide it as president
during a period of expansion and further development.

The future of the Society lies fair and bright before it. There
is every hope that it will extend its usefulness to all parts of Cali-
fornia and that its days will be days of inspiring growth and days
of true felicity. Its friends will be multiplied and it will in this,
our California, live long in the land.

NOTES ON THE GENUS RIBES IN CALIFORNIA

CLARENCE R. Quick

In the course of field work in the control of white-pine blister
rust (Cronartium ribicola Fischer) in California, field supervisors
have made interesting observations on the distribution and mor-
phology of native species of Ribes (wild currants and goose-
berries). The following notes include the more important of
these observations which have not hitherto been recorded in
botanical literature.

Acknowledgement is made of the facilities and technical
assistance furnished by the College of Agriculture, the Depart-

1938 ] QUICK: RIBES 287

ment of Botany and the Botanical Garden of the University of
California at Berkeley. The Botanical Garden has materially
assisted in accumulating and maintaining a large collection of
Ribes species, both native and exotic.

Rises amarum McClatchie, Erythea 2: 79. 1894. The north-
ern limit in the Sierra Nevada of the distribution of Ribes amarum,
a species principally of southern California, is given by Jepson
(Fl. Calif. 2: 153. 1986) as Mariposa County. Early in the
spring of 1936, however, Douglas R. Miller and Horace D. Jones
collected specimens of Ribes in a canyon northeast of George-
town, Eldorado County, which were identified by Stephen N.
Wyckoff as this species. Additional specimens were obtained by
the writer from the same locality on July 16, 1986. The collec-
tions were made at an elevation of about 3000 feet in a ravine
along that small branch of Canyon Creek which is crossed by the
Georgetown-Wentworth Springs road (section 5, T. 12 N., R. 11
E., Mt. Diablo B. & M.). During the past two years workers on
blister rust control have reported a fair abundance of the species
in this vicinity.

The elevational range of Ribes amarum is usually given as
1000 to 4000 feet. In September, 1936, Russell L. Keene showed
the writer plants of this species on Chowchilla Mountain in
Mariposa County. These plants grew on a small, steep branch
of Devil’s Gulch, at an elevation of about 6500 feet (sections 11
and 12, T. 5 S., R. 20 E., Mt. Diablo B. & M.). Specimens for

the herbarium and a few seeds were collected.

Rises cerEum Dougl. Trans. Hort. Soc. 7: 512. 1830. This
species is both widespread and abundant in the higher elevations
of the Sierra Nevada. Fruiting bushes have been observed by
the writer on the South Fork of the Stanislaus River, as low as
4800 feet, and the species has been noted by Jepson (FI. Calif. 2:
146. 1986) at 12,000 feet elevation on Mt. Whitney. The size
of the plants varies widely with elevation: at 6500 to 8500 feet,
in the Stanislaus region, whole hillsides are sometimes covered by
the species; at an elevation of about 7000 feet, the bushes may
be as tall as eight feet; while at higher elevations in the same
region where Pinus albicaulis Engelm. is slightly more than knee-
high, plants of R. cereum are but a few inches high. Several
three-inch bushes are shown in figure 1 (pl. XXXVIII), photo-
graphed at 11,000 feet elevation on a spur of Mount Leavitt near
the summit of Sonora Pass highway (Mono-Tuolumne county
line). Figure 2 (pl. XXXVIII), taken near Strawberry Lake,
Tuolumne County, at 7200 feet elevation (section 13, T. 4.N., R.
18 E., Mt. Diablo B. & M.) depicts one of the branches of fruit
from this region, which characteristically reach a greater length
in a single season than do the entire plants illustrated in figure 1.

Rises Menziesi Pursh var. ixoderme var. nov. Frutex viscido-
glandulosissimus usque ad 18 dm. altus, internodis crebre acule-

288 MADRONO [Vol. 4

atis hirsutisque, herba partium juniorium ac foliis utrinque flavido-
viridibus dense glanduloso-pubescentibus hirsutulis viscidis aro-
maticissimus, pedunculis petiolos plerumque excedentibus, floribus
amplis (14-20 mm. longis) rubiginosis, ovario pilis albidis ac
glandulis rubro-stipitatis obtecto, fructu diametro usque ad 2 cm.
aliquantulum ovoideo stramineo vel dilute sanguineo, fructus
crusta dura viscida crebre breviterque aculeata aromaticissima.
Deciduous shrub with densely glandular-hairy, viscid, pubes-
cent, yellowish-green, and strongly aromatic herbage; nodal

spines 3, unequal, about 1 cm. long; internodes densely prickly, |

very glandular and somewhat pubescent; peduncles 2—3 cm. long,
1—3-flowered, usually much longer than the petioles; flowers large
(14-20 mm. long), and showy; hypanthium 3—4 mm. long, about
as broad as long; sepals 2.5—-5 mm. wide, 6-9 mm. long, 1.25—2.5
times length of hypanthium; rich mahogany red on inside sur-
faces; petals white, about half as long as filaments; anthers usu-
ally 3 mm. or longer; ovary densely covered with red-stalked
glands, and a few white hairs; fruits subglobose or slightly
elongate, up to 2 or 2.5 cm. in diameter, yellowish, or tan-colored,
often tinged reddish, very densely glandular-bristly, viscid and
strong-smelling, the thick skins superficially nauseous because of
the oily glands.

Type: abundant in tall mixed chaparral, at 3000 feet altitude,
just below the lower limit of yellow pine along the Sand Creek
road to General Grant National Park in the foothills of the Sierra
Nevada, Fresno County (section 19, T. 14S., R. 27 E., Mt. Diablo
B. & M.), April 17, 1934, C. R. Quick 1254 (Herb. Univ. Calif. no.
575,099; isotypes, Stanford Univ., U. S. Nat. Herb., Calif. Acad.
Sci.). It was also found growing abundantly at approximately
the same elevation in Tulare County along the Orosi-Badger road
(T.15S., R. 27 E., Mt. Diablo B. & M.). Transplants and seed-
lings have been established at Berkeley in the University of Cali-
fornia Botanical Garden.

The very numerous glands on the herbage and fruits sug-
gested the varietal designation. Of the varieties of the Ribes
Menziesii aggregate, var. iroderme most closely resembles var.
leptosmum (Cov.) Jepson. It differs from typical R. Menziesi by
having (1) much greater density of hairs and stalked glands on
the herbage, (2) larger, less spiny, but much more glandular
fruit, (3) longer petioles and especially peduncles, (4) larger
flowers, (5) a much heavier and more penetrating odor. In addi-
tion, var. ivoderme grows at a higher elevation than R. Menziesu,
and was found in a location distinctly different floristically from
that previously described for R. Menziesii. The variety differs
from.R. Menziesii var. leptosmum by having (1) young twigs con-
sistently hairy, and very bristly and glandular; (2) both surfaces
of leaves densely glandular-hairy, (3) peduncles longer, (4)
flowers heavier, (5) mature fruits much lighter colored, (6) a

Beater

PLATE XXXVIII. Srupres 1x Rizes. Fig. 1. Mature bushes of Ribes
cereum at 11,000 feet elevation (note pocket knife for comparison of size).
Fig. 2. Fruiting branch of Ribes cereum at 7200 feet elevation. Fig. 3. Part
of type specimen of Ribes Menziesti var. twoderme. Fig. 4. Fruits of Ribes
Menziesti var. ivoderme. Fig. 5. Ripe fruits of Ribes Roezlii. Fig. 6.
Fruiting branches of Ribes Roezlii (Photograph by F. A. Patty).

1938] QUICK: RIBES 289

distinctive and penetrating odor. Furthermore, the two varieties
occupy distinct geographical ranges.

The geographic and taxonomic variations of the linneon Ribes
Menziesii are outlined at some length by Jepson (op. cit., p. 155-
156). The recently discovered plant treated as a variety fits
very well, it is believed, into Jepson’s understanding of the Rf.
Menziesu group. As far as can be determined, no representative
of this aggregate has been reported previously from the Sierra
Nevada.

Rises PeETIOLARE Dougl. Trans. Hort. Soc. London 7: 514.
1830. Until recently this montane species, abundant along
streams in the Pacific Northwest, had not been reported from
California. On September 18, 1936, however, George A. Root
and Hollis Day (California Ranger 89: 4. 1987), while inspect-
ing Ribes and white pines for white-pine blister rust in the Goose
Nest (Little Shasta) district of the Shasta National Forest, found
bushes of Ribes petiolare scattered for about two miles along
Shovel Creek, Siskiyou County) (section 25, T. 47 N., R. 3 W.,
Mt. Diablo B. & M.), in the ravine at 8200 feet. This site is
about nine miles south of the California-Oregon state line. Speci-
mens for the herbarium and a few seeds were collected.

In 1934 George A. Root and Robert Roney observed a few
bushes of this species at 5000 feet on Kelley Creek in the Warner
Mountains, Lake County, Oregon, only one and one-half miles
north of the California-Oregon state line (sections 17 and 18, T.
41 S., R. 21 E., Willamette B. & M.). A few days previously the
species was observed in abundance by the writer in Lake County,
Oregon, on Crane Creek (sections 7 and 8, T., 40 S., R. 21 E.).
Similar westerly flowing creeks in the Warner Mountains just
south of California-Oregon state line were carefully inspected but
the species was not found.

Rises Rorziu Regel, Gartenfl. 28: 226. 1879. The Sierra
gooseberry is the most widespread and abundant species of Ribes
in the commercial sugar pine stands of California. It ranges in
elevation from 3500 to 8500 feet in the central Sierra Nevada,
occurring locally in extraordinary abundance at 4500 to 7500
feet.

The fruits are highly variable as to size, color, shape and
spininess, those of some bushes being exceedingly spiny, while
those of others may be practically spineless. Their usual color
when ripe is dark red, but mature berries of many bushes in the
Stanislaus National Forest are bright yellow. Figure 5 (pl.
XXXVIII) shows a number of ripe fruits collected at 7200 feet
elevation in a small area near Gooseberry Camp, southeast of
Strawberry Lake, Tuolumne County (section 13, T. 4 N., R. 18
E., Mt. Dialbo B. & M.). The fourth berry from the right in
each line of fruits is fully mature, and bright yellow.

290 MADRONO [Vol. 4

In favorable years vigorous bushes of Ribes Roezlit fruit
prolifically. The plants illustrated in figure 6 (pl. XXXVIII)
were photographed at about 6300 feet elevation on Cow Creek,
a tributary of the middle fork of the Stanislaus River, Tuolumne
County (section 34, T. 5 N., R. 18 E., Mt. Diablo B. & M.).

Bureau of Entomology and Plant Quarantine,

United States Department of Agriculture,
‘Berkeley, California, November 9, 1937

A HYBRID ERIOGONUM

Hersert L. Mason

x Eriogonum Blissianum hybr. nov. (EE. giganteum Wats. x
E. arborescens Greene). Frutex compactus, hemisphericus, 9—12
dm. altus; forma et cymis E. arborescenti, bracteis E. giganteo
similis ; folia 2-10 cm. longa, oblongo-elliptica, forma inter paren-
tes intermedia, paullo revoluta, in petiolis brevibus, crassis atten-
uata, supra puberula vel glabra, subtus dense tomentosa; cymae
paniculatae; pedunculi 6-20 cm. longi; structura floralis ut in
parentibus; perianthii segmenta albida, apicibus saepe rubes-
centibus.

Compact rounded shrub, 9-12 dm. high; resembling EL. arbo-
rescens Greene in compact growth form and inflorescence, and
E. giganteum Wats. in shape and size of bracts; leaves 2-10 cm.
long, oblong-elliptic, intermediate in shape between those of the
two parent species, somewhat revolute, attenuate to a short stout
petiole; upper surface puberulent to glabrate, lower surface with
dense felt-like tomentum; cyme paniculate; peduncules 6—20 cm.
(averaging 10 cm.) long; floral structure as in both parents;
perianth segments whitish, often with pink tinge at tips.

Type: collected in Blaksley Botanic Garden, Santa Barbara,
California, July 12, 1988, M. Van Rensselaer 557 (University of
California Herbarium no. 589831).

The specimen selected as the type appeared as a volunteer in
the Eriogonum bed of the Blaksley Botanic Garden at Santa Bar-
bara, California, and similar specimens have been grown from
seed taken from E. giganteum in that garden. Thus it would
appear that E. giganteum is the female parent. As a result of
several years’ observation it can be stated that the hybrids are
probably sterile.

The widespread popularity of this Eriogonum hybrid as a
garden ornamental makes a name desirable. As an ornamental
it surpasses either parent and is highly recommended for Cali-
fornia gardens. I take pleasure in naming it in honor of the late
Anna Dorinda Blaksley Bliss, founder of the Blaksley Botanic
Garden, and her daughter Mrs. Robert Woods Bliss.

University of California,
Berkeley, August 1, 1938.

Prate XXXIX. Erioconum Buisstanum Mason.

we \
+ -
oe 2 A

1938] INDEX

291

INDEX TO VOLUME IV

New scientific names, and page numbers of principal entries of species and

varieties, are printed in bold-face type.

Abies concolor, 2; magnifica, 22, 176

Abutilon americanum, 94; incanum,
62, 78, 94; Jacquinii, 82, 94; tex-
ensis, 94; triquetrum, 94; Wrightii,
66, 94

Acacia, 120, 121, 122, 123, 124, 177,
187, 261; accola, 186; acinacea,
184, var. Latrobei, 180; acumi-
nata, 184; adunca (crassiuscula),
184; aestivalis, 184; alata, 184;
albicans, 184; amentacea, 50, 53,
pl. 55, 56, 62, 84, 91; aneura, 186;
arabica, 185; argyrolofera, 180;
argyrophylla, 185; armata, 122,
177, 178, 179, 180, 181, 182, 184,
185; aspera, 184; Baileyana, 122,
184, 185, 186, pollen of, pl. 125;
Bakeri, 186; Bancrofti, 186; bar-
bon, 179; Bartheriana, 185; Ber-
landieri, 53, 56, pl. 57, 62, 84, 91;
Berteriana, 185; Betchei, 184, 186;
bienana, 180; binervata, 185;
bispinata, 180; bispinosa, 180;
brachybotrya var. argyrophylla,
180; bynoeana, 180; caesia, 180;
calamifolia, 182; capensis, 179,
180, 185; cardiophylla, 186; Ca-
venia, 184, 185; celastrifolia, 180,
182; chordaphylla, 180; chordo-
phylla, 182; concinna, 179, 181;
confusa, 184; conspicua, 184; cor-
data, 184; cornigera, 179, 180;
corymbosa, 186; cultriformis, 122,
177, 180, 181, 182, 184, 185; cune-
ata, 183; cuspidata, 180; cyano-
phylla, 183, 185; Cyclops, 184, 185;
dealbata, 178, 179, 180, 182; de-
cipiens, 180; decora, 186; decur-
rens, 123, 178, 180, 184, 185, var.
dealbata, 177, 178, 179, 180, 181,
182, 183, 184, 185, var. Leichardtii,
186, var. mollis, 177, 181, 182, 183,
184, 185, var. normalis, 180, var.
pauciglandulosa, 186; dentifera,
184; Dietrichiana, 184, 185; dif-
fusa var. cuspidata, 180, 181; do-
doenifolia, 179, 180; dodonaefolia,
179, 180, 181, 184, 185; Douglasii,
180; elata, 122, 184, 185; elongata
var. angustifolia, 186; exsudens,
180; extensa, 186; exudens, 180;
faleata, 185, 186; falcifolia, 179,
180; farinosa, 184; Farnesiana,
50, 58, 60, 66, 84, 92, 121, 177, 179,
180, 184; flexicaulis, 111; flexi-

folia, 186; floribunda, 178, 180,
182; floribunda latifolia, 180; flori-
bunda pendula, 180; fimbriata,
184; fragrans, 184; glandulosa,
180; glauca, 185; glauca pendula,
180; glaucescens, 184, 185; glome-
rata, 185; glomerosa, 185; grandis,
180; Greggii, 92, 185; Guilfoyli-
ana, 180; hakeoides, 186; hastu-
lata, 184; heteroclada, 180; heter-
oclita, 180; heterophylla, 179, 180;
holocericea, 185; holocericena,
185; homalophylla, 182; horrida,
179, 180; Huegelii, 180; Hugelii,
180; hybrida, 180; imbricata, 182,
185; implexa, 182, 184; impressa,
180; iteaphylla, 184; ixiophylla,
180; Jonesii, 185; Julibrissin, 179,
180; juncifolia, 184; juniperina,
186; Koa, 184; latifolia, 179, 180,
182, 183, 185; La Trobay, 184; La
Trobei, 180; Latrobei, 184; lei-
ophylla, 182; leprosa, 182, 184,
185; leptoclada, 186; leptophylla,
179, 180; leucacantha, 182; leu-
cantha, 182; leucocephala, 185;
leucophylla, 180; linata, 183; line-
aris, 179, 180,,181, 182, 183; line-
ata, 182, 184, 185; linifolia, 180,
var. prominens, 182, 184, 186;
longiflora floribunda, 180; longi-
flora magnifica, 181; longifolia,
122, 177, 178, 179, 181, 182, 183,
184, 185, var. floribunda, 178, 180,
182, 183, 184, 185, var. mucronata,
181, var. latifolia, 184, var. So-
phorae, 181, 182, 184; longissima,
179, 181; lophantha, 179, 181, 185;
lunata, 185; Maideni, 185; mela-
noxylon, 122, 177, 178, 179, 181,
182, 183, 184, 185, 187; micro-
botrya, 184; mimeroides, 181;
mollissima, 123, 181, 182; mono-
phylla, 185; monospermum, 181;
montana, 181; mucronata, 181,
186; myrtifolia, 181, var. celastri-
folia, 180, 182; myrtifolia elegans,
181; neiomanii, 181; neriifolia,
122, 185, 186; nigricans, 181;
nonuttiana, 186; notabile, 186;
notabilis, 186; obliqua, 181, 184,
186; obtusata, 186; ornithoflora,
185; ovalifolia, 185; oxycedrus,
184, paludosa, 181; paradoxa, 181;
pendula, 181, 183, 184, 186, var.

292 MADRONO

glaucescens, 180; pennata, 181;
penninervis, 186; pentadenia, 122,
184; peregrina, 185; podalyriae-
folia, 123, 184, 186; polybotrya
var. foliolosa, 186; Potteri, 182;
pravissima, 184, 186; procumbens,
179, 180; prominens, 182, 184;
pruinosa, 184, 185; pubescens, 177,
181, 183; pudalosa, 181; pulchella,
122, 181, 184, var. grandis, 180,
184, var. hispidissima, 184; pul-
chella spinosa, 181; pulcherrima,
186; pycnantha, 181, 182, 184, 185;
pyracantha, 181; reclinata, 182;
Reimeri, 183; retinoides, 123, var.
floribunda, 183, 184; retinodes,
182, 184, 185, 263, var. gracillima,
184; Riceana, 183, 184, 185; rigens,
180, 182; Roemeriana, 92; rotundi-
folia, 181; rubida, 184; rubra,
184; rugata, 179, 181; rugosa, 179,
181; rutaefolia, 122; salicifolia,
179, 181; salicina, 181, var. Wayae,
184; saligna, 182, 184, 185; sentis,
185; sophora, 181; Sophorae, 182;
speciosa, 179, 181; spectabilis, 181,
185, var. excelsa, 185; stenoptera,
184; strombulifera, 179, 181; suave-
olens, 179, 181; subulata, 186;
tenuifolia, 122, 184, pollen of, pl.
125; trinerva, 179, 181; trinervata,
179, 181, 182, 185; trinervia, 181;
trinervis, 185; trineura, 179, 181,
185; undulata, 182; venulosa, 182;
verniciflua, 185; verrucosa, 182;
verticillata, 123, 177, 179, 181, 184,
187; vestita, 179, 181, 184; vir-
gata, 185; viscidula, 186

Acacias, Introduction of, into Cali-
fornia, 177

Acalypha hederacea, 93

Acanthocereus pentagonus, 58, pl. 59,
90

Acanthochiton Wrightii, 82, 90

Acanthomintha ilicifolia, 164

Acer glabrum var. Douglasii, 268

Achlys triphylla, 267

Achyranthes repens, 90

Acleisanthes Greggii, 90; obtusa, 90

Acoma, 209

Actinea linearifolia, 98; odorata, 62, 98

Additions to our knowledge of the
flora of Mount Baker, Washing-
ton, 263

Adelia Vaseyi, 56, 93

Adenocaulon bicolor, 269

Agalinis heterophylla, 78

Agarista, 209; calliopsidea, 215

Agave, 88; Lechuguilla, 64, 88; melli-
flua, pl. 65, 88; Shawii, 70

[Vol. 4

Agoseris heterophylla, 77

Agropyron trachycaulum, 265

Agrostis Thurberiana, 265

Aguapal, 53

Ailanthus glandulosa, 261, 263

Aira caryophyllea, 265

Alaska peninsula, westernmost, Flora
of the Aleutian Islands and, notice
of publication, 238

Albizzia, 120, 121, 179; Acle, 123;
fragrans, 184; Julibrissin, 123, 179,
180, 181, pollen of, pl. 125; Leb-
bek, 123, 179, 181; lophantha, 123,
179, 181, 185; occidentalis, 111

Aleutian Islands and westernmost
Alaska peninsula, Flora of, notice
of publication, 238

Algae of the Templeton Crocker expe-
dition, A preliminary report on,
notice of publication, 101

Allionia Nyctaginea, 90

Allium, 69; arenicola, 88; falcifolium,
205

Allocarya, 134

Allotropa, 137; The structure of, 137;
virgata, 137, fig. 137, pl. 143, pl.
145, pl. 147, pl. 149, pl. 151

Alsine Baldwinii, 91

Alvordia fruticosa, 110, 111

Amaranthus, 83; Berlandieri, 90; hy-
bridus, 82, 83, 90; retroflexus, 266;
spinosus, 82, 90

Amargosal, 56

Amblyolepis setigera, 98

Ambrosia bipinnatifida, 120; Chamis-
sonis, 120; elatior, 60, 83, 98;
psilostachya, 98; pumila, 165

Amelanchier alnifolia, 18; florida, 17,
18, pl. 19; florida Lindley, The
identity of, 17; humilis, 20; huron-
ensis, 20; sanguinea, 17

Ammannia coccinea, 82, 98

Ammoselinum Popei, 95

Amor, 50

Amoral, 50, 60

Amoreuxia Wrightii, 66, 94

Ampelopsis arborea, 82, 94; mexicana,
94

Amsinckia, 133

Amyris parvifolia, 52, 93

Anagallis arvensis, 262, 263

Ancistrocactus brevihamatus, 90;
Scheeri, 54, 56, 62, 64, 90

Andropogon barbinodis, 86; glomera-
tus, 86; littoralis, 80, 86; sacchar-
oides, 86

Anemone Ludoviciana, 34; Lyallii, 267

Angelica scabrida, 239

Anomalous new species of Lapsana
from China, An, 154

ih
,

1938]

Anredera vesicaria, 90

Antennaria, 133; Howellii, 269; lanata,
269; tomentella, 269

Anthemis arvensis, 269

Antigonon leptopus, 110

Aphanostephus ramosissimus, 98; skir-
robasis, 80, 98, var. Hallii, 53, 56,
66, 98

Apium Ammi, 95

Aplopappus, 74; Drummondii, 98;
phyllocephalus, 78, 80, 98

Applegate, E. I., Plants of the Lava
Beds National Monument, Cali-
fornia, review of, 237

Aquilegia scopulorum subsp. perplex-
ans, 239

Arabis, 37; glabra, 267; L., The genus,
in the Pacific Northwest, review
of, 37

Arbutus, 152; Menziesii, 280

Arceuthobium campylopodum
Tsugensis, 266

Archidendron Vaillantii, 123

Arctium Lappa, 98; minus, 269

Arctostaphylos, 152, 188; montana,
205; nevadensis, 176; patula, 176

Arenaria Douglasii, 205; macrophylla,
266; serpyllifolia, 267

Argemone alba, 91; mexicana, 91;
platyceras, 91

Aristida purpurea, 52, 53, 56, 64, 80, 86

Aristolochia longiflora, 269

Artemisia Palmeri, 165

Arundo Donax, 82, 86

Ascomycete, An unusual, in the shells
of marine animals, notice of pub-
lication, 101

Asplenium Trichomanes, 264; viride,
264

Aster exilis, 60, 98; Palmeri, 98; spi-
nosus, 98

Asteraceae from northern California,
Tracyina, a new genus of, 73

Astragalus oocarpus, 164

Astrophytum asterias, 64, 90

Atamosco texana, 58, 89

Atrichoseris platyphylla, 158

Atriplex, 78; acanthocarpa, 62, 89;
arenaria, 89; canescens, 89; mata-
morensis, 78, 80, 89

Avakian, A. M., Staff publications,
California Forest and Range Ex-
periment Station [bibliography
of] notice of publication, 102

Avena fatua, 263

Avicennia nitida, 52, 78, 80, 96

var.

Babcock, E. B., and Stebbins, G. L.,
Jr., The genus Youngia, notice of
publication, 136

INDEX 293

Baccharis, 62; glutinosa, 98; texana,
80, 98

Badilla salitrosa, 49, '7’7

Bahia absinthifolia, 98

Baja :California, map of vegetation,
fig. 109; The vegetation of the
Cape Region of, 105

Baker, M. S., An undescribed species
of Viola from Utah, 194

Baptisia leucophaea, 80, 92

Barbarea orthoceros var. dolicarpa,
267

Barettal, 62

Basicladia, 30, 32; crassa, 29

Batis maritima, 78, 80, 89

Battarea phalloides (Dicks.) Pers. in
Santa Barbara, 237

Bauer, H. L., Moisture relations in
the chaparral of the Santa Monica
Mountains, California, notice of
publication, 101

Beach pine, 285

Bedeutung der Polyploidie fiir die
Verbeitung der Angiospermen,
review of, 198

Beloperone californica, 112

Bennett juniper, 24, pl. 25, 26, 27

Berberis aquifolia, 267

Bernardia myricaefolia, 56, 93

Betula occidentalis, 266

Bidens, The genus, notice of publica-
tion, 136

Bishop pine, 285

Black mangrove, 78

Blake, S. F., Tracyina, a new genus of
Asteraceae from northern Cali-
fornia, 73

Blodgett, C. O., and Mehlquist, G. L.,
Color variation in Delphinium
cardinale Hook., 231

Bloomeria Clevelandii, 163

Boerhaavia erecta, 90; viscosa, 90

Bonar, L., An unusual ascomycete in
the shells of marine animals,
notice of publication, 101

Bordas scarp, 64

Borrichia frutescens, 78, 98

Boscaje de palma, 50, 80

Botanical studies in the Uinta Basin
of Utah and Colorado, review of,
196

Botanical survey of the Olympic Pe-
ninsula, Washington, review of,
39

Botanical terms, Lindley, Illustrated
dictionary of, review of reprint,
196

Botrychium lanceolatum, 264;
naria, 264; multifidum, 264

Bourreria sonorae, 111

Lu-

294, MADRONO

Bouteloua, 53; barbata, 52, 53, 54, 56,
62, 64, 66, 86; hirsuta, 86; trifida,
56, 62, 66, 86

Bracelin, Mrs. H. P., Ynes Mexia, 273

Brachiaria ciliatissima, 86

Brassica campestris, 267

Brickellia Coulteri, 112

Brodiaea, 132; appendiculata, 130,
131, fig. 131, 132; minor, 132; A
new Californian species of, 130;
Orcuttii, 163; stellaris, 132; syn-
andra, 132

Bromus ciliatus, 265; mollis, 263;
rigidus, 262, 263; tectorum, 265

Bryum argenteum, 85

Bumelia angustifolia, 56, 58, 62, 66, 82,
95; lycioides, 53, 58, 66, 95

Bursera, 112; laxiflora, 111; micro-
phylla, 70, pl. 107, 111, 112; odor-
ata, 111; rhoifolia, 113

Butterfield, H. M., The introduction
of acacias into California, 177

Cabbage palm, 80

Cakile americana, 78, 91; maritima
var. aequalis, 77

Calamagrostis canadensis, 265

Calceolaria verticillata, 80, 94

California, Introduction of Acacias
into, 177; Plants of Lava Bed Na-
tional Monument, review of, 237;
Early place names of, used by
Daniel Cleveland, 67

California Botanical Society, Begin-
ning years of, 276; Proceedings
of, 72, 103, 199, 239

California Forest and Range Experi-
ment Station, Staff publica-
tions, 1916-1937, [bibliography of ]
notice of publication, 102

California juniper, 22

California nutmeg tree in cultivation,
166, 167

California Salvias, review of Salvia,
section Audibertia, review of, 196

Californian species of Brodiaea, A
new, 130

Californian species of the genus Core-
opsis L., The native, 209

Calliandra, 122; californica, 111, 113;
eriophylla, 92; grandiflora, 122,
123, pollen of, pl. 125

Callirrhoé digitata, 53, 94

Callitriche palustris, 268

Calochortus Dunnii, 163; rhodothecus,
239

Calyptrocarpus vialis, 98

Camassia quamash, 266

Canary Island date palm, Hanging
gardens of the, 260

Capsella Bursa-pastoris, 267

[Vol. 4

Capsicum baccatum, 62, 97
Cardiospermum corindum, 82, 98;
Halicacabum, 52, 62, 82, 83, 93
Carex ablata, 265; arcta, 265; Brit-
toniana, 82, 88; festivella, 265;
laeviculmis, 265; leporinella, 265;
leptopeda, 265; Piperi, 265; Rossii,
265; rostrata var. utriculata, 265;
sitchensis, 265; vesicaria var.
major, 265

Carlowrightia glabrata, 97

Carpenteria, 166

Carum Gairdneri, 4

Cassia atomaria, pl. 107, 111; bauhini-
oides, 92; Medsgeri, 92; pumilio,
92

Castalia elegans, 58, pl. 59, 60, 91

Castanopsis sempervirens, 176

Castela Nicholsonii, 54, 56, pl. 57, 64,
93

Castelaria texana, 93

Cathartolinum alatum, 92; multicaule,
92

Caulanthus stenocarpus, 163

Ceanothus, 166, 199; austromontanus,
164; cyaneus, 164; Jepsoni, 205;
thyrsiflorus: Extension of range
of, 199

Celosia floribunda, 111; paniculata, 90

Celtis mississippiensis, 60, 82, 89; pal-
lida, 53, 54, 56, 58, 60, 62, 64, 80,
82, 83, 89

Cenchrus incertus, 86; pauciflorus, 62,
80, 83, 86

Cenizal, 62

Centaurea americana, 82, 98

Cerastium viscosum, 267; vulgatum,
267

Ceratophyllum demersum, 242

Cercidium, 66, 112; floridum, 64, 66,
92; peninsulare, 110, 111; sonorae,
111; texanum, 53, 66, 92

Cereus, night-blooming, 58

Cevallia sinuata, 94

Chaenactis glabriuscula, 208

Chaetomorpha, 30, 32; chelonum, 29;
A new species of, from China, 28;
sinensis, 30, pl. 31

Chamaecrista cinerea, 92; littoralis, 78,
92

Chamaesaracha conioides, 97

Chamaesyce cordifolia, 83, 93; hyper-
icifolia, 82, 93; laredana, 66, 93;
serpens, 93

Chaparral, 50, 52, 62, 64, 83, 84; black,
50

Chaparral-cenizal, 56

Chaparral-mesquital, 54

Chaparro prieto, 50

Chara praelonga, 60

Charco, 49, 60

a

1938]

Charleston Mountains, Clark County,
Nevada, Notes on the flora of,
notice of publication, 239

Charleston Mountains, Nevada, A new
Penstemon from, 128

Cheilanthes gracillima, 264; siliquosa,
205

Chenopodinae, 172

Chenopodium album,
viride, 89

Chimaphila, 146, 150, 152

China, An anomalous new species of
Lapsana from, 154; A new species
of Chaetomorpha from, 28

Chloris andropogonoides, 86; cucul-
lata, 53, 64, 86; Gayana, 86;
Petraea, 80, 86

Chorizanthe Orcuttiana, 33, 163; poly-
gonoides, 33

Chrysanthemum Leucanthemum var.
pinnatifidum, 269

Cicuta occidentalis, 268

Cienfuegosia sulphurea, 94

Cirsium undulatum, 98

Cissus incisa, 94

Citharexylum Berlandieri, 96; flabelli-

folium, 111

Cladophora, 30, 32;
nana, 29

Cladothrix lanuginosa, 78, 80, 90

Clappia suaedaefolia, 98

Clarkia delicata, 164

Claytonia, 172, 173, 174; asarifolia,
174; section Belia, 174; cheno-
podina, 171, 172, 174, pl. 175;
cordifolia, 172, 173, 174; neva-
densis, 171, 172, 173, 174, pl. 175;
nevadensis Watson, On the iden-
tity of, 171

Clematis Drummondii, 62, 82, 83, 91;
ligusticifolia, 34; Vitalba, 34

Clethra, 138, 152, 153

Cleveland, Daniel, Early California
place names used by, 67

Clokey, I. W., A new Penstemon from
the Charleston Mountains, Ne-
vada, 128; Notes on the flora of
the Charleston Mountains, Clark
County, Nevada, notice of publi-
cation, 239

Clover, E. U., Vegetational survey of
the Lower Rio Grande Valley,
Texas, 41, 77

Cocculus diversifolius, 91

Coelastrum, 124

Coelostylis texana, 93

Cogswellia, 133, 198

Coldenia canescens, 64, 66, 96

Coldenia in the Galapagos Islands,
notice of publication, 101

Collinsia parviflora, 269

54, 89, 266;

glomerata var.

INDEX

295

Collomia heterophylla, 269; linearis,
269

Cologania, 92

Color variation in Delphinium cardi-
nale Hook., 231

Colorado, Botanical studies in Uinta
Basin of Utah and, review of, 196

Colubrina glabra, 111, 113; texensis,
66, 94

Comal, 58, 66

Commelina crispa, 53, 80, 88; longi-
caulis, 60, 82, 88

Condalia obovata, 56, 94; obtusifolia,
94; spathulata, 113

Constance, L., Reviews: Applegate,
Plants of the Lava Beds National
Monument, California, 237; Gil-
key, Handbook of Northwest
flowering plants, 38; Henderson,
The early flowering of plants in
Lane County, Oregon, in 1934, 38;
Jones, A botanical survey of the
Olympic Peninsula, Washington,
39; Mathias, A revision of the
genus Lomatium, 197; Rollins,
The genus Arabis L. in the Pa-
cific Northwest, 37; St. John,
Flora of southeastern Washington
and of adjacent Idaho, 132; St.
John and Warren, The plants of
Mt. Rainier National Park, Wash-
ington, 167; Wynd, The flora of
Crater Lake National Park, 167

Constance, L., Systematic study of the
genus Eriophyllum, notice of pub-
lication, 101

Contributions from the Dudley Her-
barium, 1927-1936, notice of pub-
lication, 102

Convolvulus incanus, 53, 95

Conyza Coulteri, 98

Cooperia Drummondii, 89

Copeland, H. F., On the pollen of the
Mimosoideae and the identity of
the supposed alga Phytomorula,
120; The reproductive structures
of Pleuricospora, 1; The struc-
ture of Allotropa, 137

Corallorhiza maculata, 266

Cordia Boissieri, pl. 57, 58, 64, pl. 81,
83, 84, 96

Coreocarpus, 209

Coreopsis, 209; Bigelovii, 210, 211,
212-214, 215, 227, 228, pl. 229;
californica, 209, 210, 211, 217-220,
222, 223, 225, 228, pl. 229, 230, 231;
calliopsidea, 210, 211, 214, 215-217,
227, 228, pl. 229; var. nana, 217;
cardaminifolia, 80, 98; Distribu-
tion of native Californian species
of, fig. 210; Douglasii, 210, 211,

296

218, 219, 220-223, 228, pl. 229, 230,
231; section Euleptosyne, 211, 223,
228, 230, 231; gigantea, 210, 211,
226-227, 231; hamiltoni, 210, 211,
214-215, 227, 228, pl. 229; sub-
genus Leptosyne, 209, 228, 231; The
native Californian species of the
genus, 209, pl. 229; maritima, 209,
210, 211, 225-226, 231; section
Pugiopappus, 209, 211, 217, 227,
928, 230; Stillmanii, 209, 210, 211,
222, 223-225, 228, pl. 229, 231, var.
Jonesii, 220, 222, 223, 228; section
Tuckermannia, 209, 211, 231

Coronopus didymus, 33

Cory, V. L., Occurrence of Talinum
pulchellum in Texas, 67

Coryphantha Pirtleana, 64, 90; Run-
yonii, 64, 90

Costillal, 66

Cotoneaster, 261

Coursetia axillaris, 56, 92

Covillea, 66; tridentata, 66, 92

Cranberry from the Tahoe National
Forest, 201

Crater Lake National Park, Flora of,
review of, 167

Crepis, 156; capillaris, 260

Cristatella erosa, 91

Crockeria, 74; chrysantha, 74

Croftia parvifolia, 89

Cronartium ribicola, 286

Croton, 93; ciliato-glandulosus, 62, 80,
93; Cortesianus, 58, 93; Engel-
mannii, 93; fruticulosus, 62, 93;
leucophyllus, 83, 93; neomexi-
canus, 93; punctatus, 93; Torrey-
anus, 93

Cruciferae, Glaucocarpum,
genus in the, 232

Crum, E., Review: The California
Salvias, section Audibertia, 196

Cryptantha, 233; Grahamii, 233; nana,
233

Cryptogramma densa, 264

Cucumis Anguria, 98

Cucurbita foetidissima, 98

Cupressus Macnabiana, 263; macro-
carpa, 263; Sargentii, 205

Cuscuta arvensis, 82, 83, 95; indecora,
62, 95

Cycas revoluta, 170

Cyperus acuminatus, 60, 88; ene
latus, 60, 88; cylindricus, 88; ele-
gans, 88; erythrorrhizos, 88; ferax,
88; globosus, 88; ochraceus, 82,
88; oxycarioides, 60, 88; strigosus,
88; uniflorus, 80, 88; virens, 88

Cynodon Dactylon, 60, 66, 83, 86

Cypress, Mendocino, 285

Cyrtocarpa edulis, pl. 107, 111

a new

MADRONO

[Vol. 4

Dactylis glomerata, 265

Dactyloctenium Aegyptium, 86

Date palm, Canary Island, Hanging
gardens of, 260

Datura innoxia, 97

Daubentonia Cavinillesii, 92;
folia, 52

Daucus Carota, 268

Dayton, W. A., A cranberry from the
Tahoe National Forest, 201

Delphinium cardinale, 231; cardinale
Hook., Color variation in, 231;
nudicaule, 232; Menziesii, 267

Delta palm, 80

Desmanthus, 122; depressa, 92

Desert juniper, 22

Didymella conchae, 101

Digger pine, 27, 28

Digitalis purpurea, 269

Digitaria sanguinalis, 86

Diospyros texana, 62, 66, 82, 83, 95

Disporum oreganum, 266; Smithii, 266

Distichlis spicata, 78, 86

Ditaxis diversiflora, 239; humilis, 93

Dolicholus americanus, 80

Dolicothele sphaerica, 53, 56, 64, 90

Dondia, 89; conferta, 89; linearis, 89;
multiflora, 78, 89

Draba prealta, 267; stenoloba, 267

Drosanthe, 272, 273

longi-

Dryopteris, 261; arguta, 262, 263;
Phegopteris, 264
Dudley Herbarium, Contributions

from, 1927-1936, notice of publi-
cation, 102
Dulichium arundinaceum, 266
Dyssodia Berlandieri, 98; ; tephroleuca,
66, 98

Early California place names used by
Daniel Cleveland, 67

Early flowering of plants in Lane
County, Oregon, in 1934, review
of, 38

Eastwood, A., and Howell, J. T.,
Leaflets of western botany, notice
of publication, 102

Ebanal, 58

Echinocereus angusticeps, 53, 90;
Blanckii, 90; enneacanthus, 56,
62, 64, 66, 90; Fitchii, 64, 90;
papillosus, 53, 56, 90; pentalophus,
54, 56, 64, 90 |

Echinochloa colonum, 82, 86; Crus-
galli, 82, 86; Crusgalli zelayensis,
86

Echinocystis fabacea, 263

Kchinodorus cordifolius, 60, 86

Echium, 33; plantagineum, 34

Eclipta alba, 82, 98

Ehretia elliptica, 96

1938]

Eichornia crassipes, 60, 88

Eleocharis albida, 78, 80, 88; obtusa,
266; palustris, 88

Elephant tree, 70

Eleusine indica, 86

Elymus glaucus, 265

Encelia farinosa, pl. 107, 113

Entada, 122

Enterolobium, 122

Ephedra antisiphilitica,
pedunculata, 86

Epigaea, 9

Epilobium minutum, 205, 268; panicu-
latum, 268

Epling, C., The California Salvias:
review of Salvia, section Audi-
bertia, review of, 196; Notes on
Stachys rigida Nutt., 270

Equisetum hyemale, 264; limnosum,
264

Eragrostis Barrelieri, 86; Beyrichii,
87; curtipedicellata, 53, 64, 87;
hypnoides, 82, 87; lugens, 87; rep-
tans, 87; secundiflora, 53, 80, 87;
sessilispica, 80, 87; spicata, 87

Eremocarpus Bentham: Preoccupied?
272

Eremodaucus, 273

Eremosporus, 272;
theae, 273

Erigeron acris var. debilis, 269; an-
nuus, 269; argentatus, 233; cana-
densis, 98, 270; philadelphicus,
270; repens, 78, 98; tenuis, 82, 98

Eriobotrya japonica, 262, 263

Eriochloa punctata, 87

Eriogonum, 208; Abertianum, 189,
190; Abertianum and its varie-
ties, 189, var. bracteatum, 190,
192, 193, var. cyclosepalum, 190,
192, var. Gillespiei, 190, 193, var.
lappulaceum, 190, 193, subsp. lap-
pulaceum, 190, 193, var. neomexi-
canum, 189, 190, 191, var. ruber-
rimum, 189, 190, 191, subsp. typi-
cum, 190, var. Villosum, 190, 191,
192; arborescens, 290; arizonicum,
189; Blissianum, pl. facing 290,
290; cyclosepalum, 189, 192; gi-
ganteum, 290; A hybrid, 290, pl.
facing 290; lappulaceum, 193;
multiflorum, 89; ovalifolium, 198;
pharnaceoides, 189; pinetorum,
189, 191; vimineum var. caninum,
205

Eriophyllum, 101; lanatum, 270; A
systematic study of the genus,
notice of publication, 101

Erodium cicutarium, 268

Erpodium domingense, 85

Eryngium compactum, 95

62, 64, 86;

section Drosan-

INDEX

297

Erysimum cheiranthoides, 267
Erythrina herbacea, 82, 92

Escobaria Runyonii, 90

Esenbeckia flava, 111; Runyonii, 93
Ksteros, 48

Ethnography, Tiibatulabal, review of,

236

Eucalyptus, 261

Eugenia Hookeri, Seedlings from
polyembryonic seeds of, 115, pl.
117

Eupatorium ageratifolium, 98; azur-
eum, 98; odoratum, 80, 82, 98;
sagitatum, 110

Euphorbia tomentulosa, 113; Xanti,
111

Eurytaenia texana, 95

Eustoma Russellianum, 95

Evax multicaulis, 98

Evolvulus alsinoides, 95

Eysenhardtia texana, 92

Ferocactus hamatacanthus, 54, 66, pl.
79, 90

Festuca megalura, 262, 263; myuros,
263; rubra, 265; supina, 265; sub-
ulata, 265

Fimbristylis castanea, 78, 88

Fir, red, 22; silver-tip, 2

Flora of the Aleutian Islands and

westernmost Alaska peninsula
with notes on the flora of the
Commander Islands, notice of

publication, 238

Flora of Crater Lake National Park,
review of, 167

Flora of Mount Baker, Washington,
Additions to our knowledge of,
263

Flora of San Diego County, Califor-
nia, Notes on, 33

Flora of southeastern Washington
and of adjacent Idaho, review of,
132

Forestiera angustifolia, 53, 54, 58, 80,
82, 95

Florestina tripteris, 98

Flowering plants, Handbook of North-
west, review of, 38

Flowering of Wolffiella
(Hegelm.) Hegelm., 241

Fouquieria peninsularis, 111, 113

Fosberg, F. R., Eriogonum Abertia-
num and its varieties, 189

Fragaria vesca, 267

Franseria ambrosioides, 113; bipinna-
tifida, 119, 120; bipinnatifida du-
bia, 120; bipinnatifida villosa, 120;
Chamissonis, 119, 120, subsp. bi-
pinnatisecta, 120, var. bipinnati-
secta, 120, var. cuneifolia, 120,

lingulata

298 MADRONO

var. mialvaefolia, 120, subsp.
typica, 120; confertiflora, 80, 98;
cuneifolia, 120; Lessingii, 120;
magdalenae, 113; Maritime, of the
Pacific Coast, 119; villosa, 120

Fraxinus Berlandieri, 60, 82, 95

Fremontia, 166

Fritillaria lanceolata, 266

Froelichia campestris,
mondii, 90

Frullania eboracensis, 85; squarrosa,
85

Fucus, 127

90; Drum-

Gaertneria Chamissonis, 120; bipin-
natifida, 120; bipinnatifida dubia,
120

Gaillardia pulchella, 53, 78, 80, 98

Galapagos Islands, The plant genus
Coldenia in, notice of publication,
101

Galarhoeus, 93; arkansanus, 93

Galeopsis Tetrahit, 269

Galium Aparine, 97, 269; bifolium,
269; Claytoni, 269; verum, 36

Galpinsia Hartwegii, 95; tubicula, 80,
95

Gander, F. F., Notes on some San
Diego County endemics, 163;
Notes on the flora of San Diego
County, California, 33

Gardner, Nathaniel Lyon, 126, pl.
facing 126; A new species of
Chaetomorpha from China, 28

Gardner, N. L. and Setchell, W. A.,
Iridophycus, with special refer-
ence to the South American
species, notice of publication, 101

Gaultheria humifusa, 268

Gaura Drummondii, 95; parviflora,
82; sinuata, 34; villosa, 95

Gayoides crispum, 62, 94

Gentiana Amarella, 269

Genus Arabis L. in the Pacific North-
west, review of, 37

Genus Bidens, notice of publication,
136

Genus Youngia, notice of publication,
136

Geraea viscida, 165

Geranium carolinianum, 92; molle,
268

Gerardia heterophylla, 97

Gilia incisa, 58, 96; latifolia, 158;
polycladon, 233

Gilkey, H. M., Handbook of North-
west flowering plants, review of,
38

Ginkgo biloba, 170

Glaucocarpum, 233, 234; A new genus

[Vol. 4

in the Cruciferae, 232; suffru-
tescens, 233, pl. 235

Glock, W. S., Observations on the
western juniper, 21

Glyceria elata, 265

Gnaphalium purpureum, 270; spa-
thulatum, 98; uliginosum, 270

Gobernadoral, 66

Gochnatia arborescens, pl. 107, 111;
hypoleuca, 98

Goniostachyum citrosum, 62, 96

Gooseberry, Sierra, 289

Gossypium Davidsonii, 111

Graham, E. H., Botanical studies in
the Uinta Basin of Utah and
Colorado, review of, 196

Grass, coastal marsh, 49

Grasses of Montana, A key to, based
upon vegetative characters, notice
of publication, 101

Grindelia colepsis, 98

Guaiacum sanctum, 56, 92

Gutierrezia eriocarpa, 98

Habenaria dilatata, 266

Haddock, P. G., Picea Breweriana in
Shasta County, 176

Haematoxylon brasiletto, 111

Hamatocactus setispinus, 54, 56, 58,
62, 64, 90

Hamosa Nuttalliana, 80, 92

Handbook of Northwest flowering .

plants, review of, 38
Hanging gardens of _ the
Island date palm, 260

Haplocladium microphyllum, 85

Hartmannia speciosa, 60, 95

Hartwell, R., Battarea phalloides
(Dicks.) Pers. in Santa Barbara,
237

Hechtia, 64; texensis, 64, pl. 65, 84,
88

Heimia salicifolia, 58, 98

Helenium linifolium, 98; microceph-
alum, 98; quadridentatum, 98

Helianthus annuus, 99; cucumerifolius,
99

Helietta parvifolia, 52, 62, pl. 63, 93

Heliotropium confertifolium, 62, 66,
96; curassavicum, 96; indicum,
58, 96; inundatum, 96; parvi-
florum, 96

Helleranthus quadrangulatus, 96

Henderson, L. F., The early flowering
of plants in Lane County, Ore-
gon, in 1934, review of, 38

Heteranthera limosa, 60, 88

Heteropogon contortus, 87

Heterotheca subaxillaris, 99

Heuchera brevistaminea, 163

Canary

eed aaa

1938]

Hibiscus cardiophyllus, 94

Hippurus vulgaris, 268

Hitchcock, C. L., A key to the grasses
of Montana based upon vegetative
characters, notice of publication,
101

Hoffmannseggia caudata, 92; densi-
flora, 78, 92

Holcus halepensis, 60, 83, 87; lanatus,
265

Homalocephala texensis, 56, 90

Hookera, 132

Hoover, R. F., A new Californian
species of Brodiaea, 130

Hordeum murinum, 263

Howell, J. T., The plant genus Col-
denia in the Galapagos Islands,
notice of publication, 101

Howell, J. T., and Eastwood, A.,
Leaflets of western botany, notice
of publication, 102

Huisachal, 50, 60

Huisache, 50

Hultén, E., Flora of the Aleutian
Islands and westernmost Alaska
peninsula with notes on the flora
of the Commander Islands, notice
of publication, 238

Hybrid Eriogonum, 290

Hydrocotyle umbellata, 95

Hymenopappus lugens, 233

Hypericophyllum, 154

Hypochaeris radicata, 270

Hypopitys, 3, 9, 10, 11, 152

Hyptis tephrodes, 111

Ibervillea Lindheimeri, 64, 98

Idaho, Flora of southeastern Wash-
ington and adjacent, review of,
132

Identity of Amelanchier florida Lind-
ley, 17

Indigofera leptosepala, 78, 80, 92

Iliamna Greene, A resurrection and
revision of the genus, notice of
publication, 102

Illustrated dictionary of botanical
terms, Lindley, review of reprint,
196

Inga, 121, 122; anomala, 121, 123;
cordistipula, 123; edulis, 123;
myriantha, 123; spectabilis, 123;
tergemina, 122, 123

Introduction of acacias
fornia, 177

Inyo County, California, check list of
plants of Rock Creek Lake Basin,
notice of publication, 239

Ionoxalis, 82, 92; violacea, 92

Ipomoea carnosa, 95; dissecta, 95;
fistulosa, 50, 60, 95; Pes-Caprae,

into Cali-

INDEX

299

78, 95; sinuata, 96; stolonifera,
78; trifida, 96
Iridophycus, 127; with special refer-
ence to the South American
species, notice of publication, 101
Isoetes echinospora var. Braunii, 265
Iva angustifolia, 99; ciliata, 99
Ixeris, 154, 157

Jatropha Berlandieri, 66, 93; cinerea,
110, 111, 113; spathulata, 53, 56,
62, 64, 93, pl. 107, 113; stimulosa,

93

)

Jeffrey pine, 22, 27, 169

Jepson, W. L., frontispiece; Flora of
California, notice of publication,
70; Viae felicitatis: the beginning
years of the California Botanical
Society, 276

Johnson, A. M., Seedlings from poly-
embryonic seeds of Eugenia
Hookeri, 115

Jones, G. N., A botanical survey of
the Olympic Peninsula, Washing-
ton, review of, 39

Juglans regia, 280

Juncus aristulatus, 88; bufonius, 266;
macer, 266

Juniper, Bennett, 24, 26, 27; Cali-
fornia, 22; desert, 22; western, 21,

* 22, 23, 26; Observations on, 21

Juniperus californicus, 22; occiden-
talis, 21, pl. 25; utahensis, 22

Jussiaea californica, 33, 242; diffusa,
60, 95

Justicia Runyonii, 97

Kallstroemia parviflora, 80, 92

Karwinskia Humboldtiana, 56, 62, 64,
94, 111

Keck, D. D., Studies in Penstemon
VI. The section Aurator, notice
of publication, 239

Key to the grasses of Montana based
upon vegetative characters, notice
of publication, 101

Koeberlinia spinosa, 64, 94

Kosteletskya hastata, 94

Kneiffia arenicola, 95

Lactuca, 154; scariola var. integrata,
270; spicata, 270

Lantana horrida, 52, 54, 58, 82, 96;
macropoda, 96

Lapsana, 154, 157; An anomalous new
species of, from China, 154; apo-
gonoides, 154, pl. 155, 156; com-
munis, pl. 155, 157; humilis, pl.
155, 156; uncinata, 154, pl. 155,
156, 157

Larkspur, scarlet, 232

300 MADRONO

Larrea, 112

Lasthenia, 74; glabrata, 74

Lathyrus palustris var. myrtifolius,
268

Lava Beds National Monument, Cali-
fornia, Plants of, review of, 237

Leaflets of western botany, notice of
publication, 102

Ledum, 10; glandulosum, 36, 176;
groenlandicum, 36

Lemaireocereus, 112; Thurberi, pl.
107, 110, 111, 112

Lemna, 241, 242, 246, 248, 250; cyclo-
stasia, 242; minor, 60, 88, 242

Lenophyllum texanum, 91

Lepachys columnaris var. pulcherrima,
66, 99

Lepidium austrinum, 91; medium,
267; virginicum, 60, 80, 91

Leptoloma cognatum, 87

Leptochloa dubia, 87; filiformis, 87;
Nealleyi, 87; uninervia, 87

Leptosyne, 209, 228, 231; Bigelovii,
212; californica, 217, 220; calliop-
sidea, 215, var. nana, 215; Doug-
lasii, 217, 218, 219, 220, 222; gi-
gantea, 226; hamiltonii, 214, 215;
Newberryi, 217, 220; maritima,
225; Stillmanii, 223

Lesquerella auriculata, 91; densiflora,
91; lasiocarpa, 91

Lessingia germanorum, 208

Leucaena glauca, 122, 180, 185; micro-
carpa, 111; pulverulenta, 82, 92

Leucophyllum, 62; frutescens, 53, 54,
pl. 55, 56, pl. 57, 58; 62,.84,) 97,
f. albiflorum, 97; minus, 97

Liagora, 71

Limnia, 174

Limonium Nashii, 78, 95

Linanthus, 157; Bigelovii, 159; Bo-
landeri, 208; dichotomus, 159;
Harknessii, 160, pl. 161; Jonesii,
158, 159, 160, pl. 161; mohavensis,
158, 159, 160, pl. 161; pharna-
ceoides, 160; septentrionalis, 159,
160, pl. 161; Two new species of,
from western North America, 157

Linaria canadensis, 97

Lindley, J., Illustrated dictionary of
botanical terms, review of reprint,
196

Linum leptopoda, 233

Lippia alba, 82, 96; Berlandieri, 96;
formosa, 111; lingustrina, 54, 58,
64, 96; macrostachya, 64, 96

Lithocarpus densiflora, 137

Lithospermum matamorense, 96

Llano salitroso, 49

Llanos, 49

Lloydia serotina, 266

[Vol. 4

Lobelia brachypoda, 98; Cliffortiana,
98; Feayana, 98

Lodgepole pine, 22, 27

Lomatia, 198

Lomatium, 198; A_ revision of the
genus, review of, 197

Lomeria, 50

Lonicera ciliosa, 269; utahensis, 269

Lophophora Williamsii, 64, pl. 79, 90

Lupinorum, Species, review of, 236

Lupinus, 133, 236, 282; albifrons
var. Brandegeei, 237; latifolius
var. Wigginsii, 237; Lyallii, 268;
Mearnsii, 237; texensis, 53, 92

Luzula campestre, 266

Lycium Berlandieri, 97; carolinianum,
54, 80, 84, 97; Chateaui, 97; hali-
mifolium, 34; Torreyi, 97

Lycopersicon cerasiforme, 60, 97

Lycopodium annotinum, 264; ob-
scurum, 264; Selago, 264

Lycopus uniflorus, 269

Lygodesmia texana, 99

Lysiloma candida, 110, 111; micro-
phylla, 110, 111

Lysimachia thyrsiflora, 268

Lythrum alatum, 98

Machaerocereus gummosus, 110, 111,
113

Macrosiphonia Macrosiphon, 97

Madia glomerata, 270

Mahonia, 166

Malacothrix obtusa, 208

Malus fusca, 267

Manfreda maculosa, 89; variegata, 89

Mangrove, black, 78

Malachra urens, 82, 94

Malpighia glabra, 52, 62, 82, 93

Malvastrum coromandelianum, 58, 94;
spicatum, 82, 94

Malvaviscus Drummondii, 82, 94

Marilaunidium hispidum, 62, 96;
jamaicense, 96; macranthum, 96

Maritime Franseria of the Pacific
Coast, 119

Marsilea macropoda, 60, 86; vestita,
60, 86

Martynia fragrans, 97

Mason, H. L., The flowering of Wolf-
fiella lingulata (Hegelm.) He-
gelm., 241; Harry Stanley Yates,
187; A hybrid Eriogonum, 290;
Two new species of Linanthus
from western North America,
157; Review, Presnall and Pa-
traw, Plants of Zion National
Park, 198; Review, Smith, Species
Lupinorum, 236; Review, Voe-
gelin, Tiibatulabal ethnography,
236

1938]

Mathias, M., Review, Graham, Botani-
cal studies in the Uinta Basin of
Utah and Colorado, 196; reprint
of Lindley, illustrated dictionary
of botanical terms, 196; A revi-
sion of the genus Lomatium,
review of, 197

Matricaria suaveolens, 270

Maytenus phyllanthoides, 93

McMinn, H. E., Ceanothus thyrsi-
florus: Extension of range, 199

Medanos, 77

Medicago Lupulina, 268

Mehlquist, G. L., and Blodgett, C. O.,
Color variation in Delphinium
cardinale Hook., 231

Melampodium cinereum, 99

Melochia pyramidata, 94; tomentosa,
111

Melosmon cubense, 96

Melothria pendula, 82, 98

Mendocino cypress, 285

Menodora heterophylla, 64, 95

Mentha arvensis var. canadensis, 269,
var. glabrata, 269

Mentzelia, 233; micrantha, 33

Menyanthes trifoliata, 269

Mesquital, 49, 52, 53, 54, 58, 83

Mesquital-chaparral, 54, 56, 58, 64,
66, 80

Mesquital climax, 52

Mesquital-nopalera, 53, 54

Mesquital-zacatal, 52, 53, 83

Metastelma barbigerum, 82, 95

Mexia, Ynes, 273, fig. 274

Mexianthus mexicanus, 274

Microsteris gracilis, 269

Micromeria pilosiuscula, 96

Microrhamnus ericoides, 62, 66, 94

Mimosa, 121, 122; Berlandieri, 82, 92;
Brandegei, 111; pudica, 122;
strigillosa, 50, 78, 92; Xanti, 111

Mimosal, 50

Mimosoideae, pollen of, and identity
of the supposed alga Phytomo-
rula, 120

Mimulus Bolanderi, 208;
269; moschatus, 269

Mirov, N. T., Phylogenetic relations
of Pinus Jeffreyi and Pinus
ponderosa, 169

Mohavea brevifolia, 158

Moisture relations in the chaparral of
the Santa Monica Mountains,
California, notice of publication,
101

Mollugo verticillata, 90

Monanthochloe littoralis, 54, 78, 80, 87

Monarda, 96; dispersa, 62, pl. 63, 96;
punctata, 96

guttatus,

INDEX

301

Monardella lanata, 165

Moneses uniflora, 268

Monoptilon bellioides, 74; bellidiforme,
74

Monotropa, 8, 10, 11, 138, 152, 153;
uniflora, 8, 268

Monotropastrum, 11

Monoxalis dichondraefolia, 92

Montana, A key to the grasses of,
based upon vegetative characters,
notice of publication, 101

Monterey pine, 26

Montia, 172, 173, 174; alpina, 173, 174;
asarifolia, 172, 174; californica,
174; fontana, 173; mnevadensis,
173, 174; perfoliata, 262, 263

Morrison, J. L., Studies in the genus
Streptanthus Nutt. I. Two new
species in the section Euclisia
Nutt., 204

Mortonia Greggii, 56, pl. 57, 84, 93

Mount Baker, Washington, Addi-
tions to our knowledge of the
flora of, 263

Mount Rainier National Park, Wash-
ington, Plants of, review of, 167

Mount Rainier, Washington, A new
Thalictrum from, 114

Muenscher, W. C., Additions to our
knowledge of the flora of Mount
Baker, Washington, 263

Muhlenbergia filiformis, 265

Myriophyllum exalbescens, 33; hip-
purioides, 242

Nacahuital, 58

Naias guadalupensis, 60, 86

Native Californian species of the
genus Coreopsis L., 209

Nemophila parviflora, 269

Neomammillaria applanata, 90; hemi-
sphaerica, 53, 54, 56, 62, 64, 90;
Heyderi, 56, 64, 90; multiceps, 90

Neowashingtonia robusta, 80

Nepeta hederacea, 269

Nerisyrenia camporum, 91

Nevada, Notes on the Flora of the
Charleston Mountains, Clark
County, notice of publication, 239

New Californian species of Brodiaea,
130

New Penstemon from the Charleston
Mountains, Nevada, 128

New records of vascular plants in
Washington, 34

New Thalictrum from Mount Rainier,
Washington, 114

New species of Chaetomorpha from
China, 28

Newberrya, 3, 5, 11, 146, 152, 153

302

Nicotiana, 71; attenuata, 236; Bige-
lovii, 236; glauca, 97; longiflora,
66; repanda, 62, 97

Nielsen, E. L., The identity of Ame-
lanchier florida Lindley, 17

Night-blooming cereus, 58

Nopal, 50

Nopalera, 50

North American species of Rumex,
notice of publication, 136

Notes and news, 69, 101, 134, 168,
199, 237, 291

Notes on the flora of the Charleston
Mountains, Clark County, Ne-
vada, notice of publication, 239

Notes on the flora of San Diego
County, California, 33

Notes on the genus Ribes in Cali-
fornia, 286

Notes on some San Diego County
endemics, 163

Notes on Stachys rigida Nutt., 270

Notholaena dealbata, 86

Nyctaginia capitata, 90

Observations on the western juniper,
21

Occurrence of Talinum pulchellum in
Texas, 67

Oenothera Drummondii, 78, 95; lacini-
ata var. mexicana, 53

Oenanthe sarmentosa, 268

Olympic Peninsula, Washington, A
botanical survey of, review of,
39

On the identity of Claytonia neva-
densis Watson, 171

On the pollen of the Mimosoideae and
the identity of the supposed alga
Phytomorula, 120

Oplismenus setarius, 82, 87

Opuntia, 50, 54, 80; Allairei, 78, 80,
91; cholla, pl. 107, 110, 111, 112;
Engelmannii, 91; fuliginosa, 111;
leptocaulis, 53, 56, 58, 62, 64, 91;
Lindheimeri, 53, 56, 58, 62, 64,
83, 91; Schottii, 56, 91

Oregon, The early flowering of plants
in Lane County, in 1934, review
of, 38

Orobanche uniflora, 269

Osmorrhiza obtusa, 268

Othake robustum, 80, 99; texanum, 99

Oxalis corniculata, 92

Oxycoccus, 202; intermedius, 202;
macrocarpos, 201, 202, pl. 203;
oxycoccos, 202; oxycoccos inter-
medius, 202; palustris, 201, 202,
var. intermedius, 202

Oxymitra androgyna, 85

MADRONO

[Vol. 4

Pachycereus, 112; pecten-aboriginum,
pl. 107, 110, 111; Pringlei, 112
Paeonia, 252; albiflora, 258; Brownii,
252, 253, 254, 255, 256, 257, 258,
pl. 259; californica, 252, 253, 254,
255, 256, 257, 258, pl. 259; Dela-
vayi, 257, 258; subgenus Moutan,
257; subgenus Onaepia, 257; sub-
genus Paeon, 257; The western
American species of, 252, pl. 259

Palm, cabbage, 80; delta, 80

Palma, boscaje de, 50, 80

Palma de micharos, 80

Palo verde, 66

Panicum fasciculatum, 82, 87; firmu-
lum, 83, 87; hians, 87; Hallii, 56,
64, 66, 83, 87; nodatum, 87; pur-
purascens, 82, 87; sphaerocarpon,
87; texanum, 87

Pantano, 49

Pappophorum bicolor, 64,
mucronulatum, 62, 66, 87

Parietaria obtusa, 89

Parkinsonia aculeata, 49, 50, 58, pl.
59, 66, 84, 92

Parosela aurea, 92; humilis, 92; nana,
62, 66, 92; pogonathera, 92

Parthenium Hysterophorus, 53, 58, 60,
66, 82, 83, 99

Paspalum conjugatum, 82, 87; dila-
tatum, 87; distichum, 78, 87;
Langei, 60, 83, 87; lividum, 87;
stramineum, 87; Urvillei, 87

Passiflora foetida, 94; lutea, 82, 94

Pastinaca sativa, 268

Patraw, P. M., and Presnall, C.,
Plants of Zion National Park,
review of, 198

Pedilanthus macrocarpus, 113

Peirson, F. W., Plants of Rock Creek
Lake Basin, Inyo County, Cali-
fornia, a check list, notice of pub-
lication, 239

Pentachaeta, 73, 74; aurea, 73, 74

Penstemon, 130; eriantherus_ var.
grandis, 239, var. redactus, 239;
fruticosus, 134; subgenus Glabri,
130; Jamesii subsp. breviculus,
239; Keckii, 128, 129, 130; A
New, from the Charleston Moun-

66, 87;

tains, Nevada, 128; speciosus,
130; Studies in VI, _ section
Aurator, notice of publication,

239; Whitedii subsp. tristis, 239
Perezia runcinata, 99
Perityle microglossa, 99
Persicaria longistyla, 82, 89; mexicana,
82, 89; pennsylvanica, 89
Petalostemon emarginatus, 78, 92
Petunia parviflora, 97

1938]

Phacelia pachyphylla, 158; parviflora,
96; patuliflora, 82, 96

Phalaris caroliniana, 87

Phaulothamnus spinescens, 90

Philadelphus Gordonianus, 267

Philoxerus vermicularis, 78, 90

Phleum pratense, 265

Phoenix canariensis, 263

Phoradendron flavescens, 53, 89

Photinia, 261

Phlox pilosa var. detonsa, 96

Phragmites communis, 60, 87

Phyla nodiflora, 96

Phylogenetic relations of Pinus Jef-
freyi and Pinus ponderosa, 169

Phymatotrichum omnivorum, 44

Physcomitrium immersum, 85;
binatum, 85

Physalis Carpenteri, 97; mollis, 80,
. 97; viscosa, 80, 97, var. spathulae-
folia, 97

Physocarpus opulifolius, 267

Phytomorula, 124; regularis, 124;
supposed alga, identity of, and
pollen of Mimosoideae, 120

Picea, 176; Breweriana in Shasta
County, 176; sitchensis, 265

Pine, beach, 285; bishop, 285; Digger,
27, 28; Jeffrey, 22, 27, 169; lodge-
pole, 22, 27; Monterey, 26; pon-
derosa, 169; western yellow, 22,
26, 27, 169

Pinus albicaulis, 287; aristata, 129;
contorta, 265; Jeffreyi, 22, 169,
170, 171, 176, and Pinus ponder-
osa, Phylogenetic relations of,
169; Lambertiana, 176; mono-
phylla, 170; murrayana, 22; pon-
derosa, 22, 169, 170, 171, and
Pinus Jeffreyi, Phylogenetic rela-
tions of, 169; radiata, 26, 170, 262,
263; Sabiniana, 27, 170, 262, 263;
scopulorum, 129

Piptadenia peregrina, 185

Piscaria, 272

Pithecolobium, 121, 122; albicans, 184;
brevifolium, 58, 82, 92; confine,
111; dulce, 123; flexicaulis, 186;
tortum, 111

Pityopus, 3, 5, 11, 146, 148, 152

Plant genus Coldenia in the Galapagos
Islands, notice of publication, 101

Plants of Mt. Rainier National Park,
Washington, review of, 167

Plants of Rock Creek Lake Basin,
Inyo County, a check list, notice
of publication, 239

Plants of the Lava Beds National
Monument, California, review of,
237

tur-

INDEX

303

Plants of Zion National Park, review

of, 198
Plantago lanatifolia, 97; lanceolata,
269; rhodosperma, 80, 97; vir-

ginica var. longifolia, 97

Platanus, 262

Plectritis, 238

Pleuricospora, 1, 140, 146, 150, 152,
153; densa, 1; fimbriolata, 1, fig.
2, pl. 12, pl. 13, pl. 14, pl. 15, pl.
16; longipetala, 1; The reproduc-
tive structures of, 1

Pluchea camphorata, 80, 99

Plumbago scandens, 62, 82, 95

Plumeria acutifolia, 111

Pital, 66

Poa alpina, 265; annua, 265; arctica,
265; epilis, 265; gracillima, 265;
palustris, 265; pratensis, 265;
secunda, 265

Fogogyne Abramsii, 165; nudiuscula,
16

Poinciana Gillesii, 179

Poinsettia heterophylla, 82, 93

Podostemma, 78, 95; longicornu, 95

Polanisia graveolens, 35; trachy-
sperma, 35, 91

Pollen of Mimosoideae, and identity

of the supposed alga Phyto-
morula, 120
Polyembryonic seeds of Eugenia

Hookeri, Seedlings from, 115
Polygala alba, 80, 93; Boykinii, 93;
macradenia, 93
Polygonum aviculare, 266; Convol-
vulus, 266; Douglasii, 266; Hydro-
piper, 266; Persicaria, 266; vivi- ©
parum, 266
Polyploidie, Die Bedeutung der, fiir
die Verbreitung der Angiosper-
men, erléutert an den Arten
Schleswig-Holsteins, mit Aus-
blicken auf andere Florengebiets,
review of, 198
Polypodium, 261;
263
Polypogon monspeliensis, 80, 87
Polystichum Lemmonii, 264
Ponderosa pine, 169
Populus tremuloides, 266; trichocarpa,
266
Porlieria angustifolia, 62, 64, 93
Portulaca oleracea, 78, 91; pilosa, 77
Potamogeton americanus, 265; epi-
hydrus, 265; tenuifolius, 265
Potentilla cryptocaulis, 239; glauco-
phylla, 267; norvegica var. hir-
suta, 268; palustris, 268 —
Preliminary report on the algae of the
Templeton Crocker’ expedition,
notice of publication, 101

californicum, 262,

304

Presnall, C., and Patraw, P. M.,
Plants of Zion National Park,
review of, 198

Proceedings of the California Botani-
cal Society, 72, 103, 199, 239

Prosopis, 122; glandulosa, 111, 180;
juliflora var. glandulosa, 49, 52,
53, 54, 56, 58, 64, 80, 83, 92; Pal-
meri, 111; strombulifera, 179, 181

Prunus avium, 268; communis, 263;
ilicifolia, 263

Psilostrophe gnaphalodes, 99

Pteridium aquilinum, 264

Pterospora, 1, 11, 138, 146, 148, pl.
149, 150, 152

Pterygoneurum cavifolium, 85

Pugiopappus Bigelovii, 212; Breweri,
212; calliopsidea, 215

Pyracantha, 261, 263

Pyramidula tetragona, 85

Pyrola, 10, 138, 146, 150, 152, 153;
chlorantha, 268

Pyrrhopappus grandiflora, 99; multi-
caulis, 99

Quercus agrifolia, 262, 263; densiflora,
137; durata, 205; virginiana, 89

Quick, C. R., Notes on the genus
Ribes in California, 286

Quincula lobata, 97

Randia aculeata, 54, 56, 97; armata,
111; Thurberi, 111

Radicula Walteri, 60, 91

Ranunculus acris, 267; aquatilis var.
capillaceus, 267; Bongardii, 267;

Macounii, 267; orthorhynchus,
267; repens, 267; reptans var.
ovalis, 267

Ravenelia, 238

Rechinger, K. H. Jr., The North
American Species of Rumex,

notice of publication, 136

Red fir, 22

Reproductive structures of Pleurico-
spora, 1

Resacas, 48, 49, 50, 84

Reseda alba, 34

Resurrection. and revision of the genus
Iliamna Greene, notice of publica-
tion, 102

Retama, 49, 58

Retamal, 49, 58

Reviews: Epling, The California Sal-
vias. A review of Salvia, sec-
tion Audibertia, 196; Gilkey,
Handbook of Northwest flowering
plants, 38; Graham, Botanical
studies in the Uinta Basin of
Utah and Colorado, 196; Hender-
son, The early flowering of plants

MADRONO

[Vol. 4

in Lane County, Oregon, in 1934,
38; Jones, A botanical survey of
the Olympic Peninsula, Washing-
ton, 39; reprint of Lindley, Illus-
trated dictionary of botanical
terms, 196; Mathias, A revision
of the genus Lomatium, 197; Pres-
nall and Patraw, Plants of Zion
National Park, 198; Rollins, The
genus Arabis L. in the Pacific
Northwest, 37; St. John, Flora of
southeastern Washington and of
adjacent Idaho, 132; St. John and

Warren, The plants of Mount
Rainier National Park, Wash-
ington, 167; Tischler, Die Be-

deutung der Polyploidie fiir die
Verbreitung der Angiospermen,
erliutert an den Arten Schleswig-
Holsteins, mit Ausblicken auf
andere Florengebiets, 198; Wynd,
The flora of Crater Lake National
Park, 167

Revision of the genus
review of, 197

Rhamnus californica, 279; Purshiana,
279

Rhizoclonium, 30, 32

Rhododendron, 10; albiflorum, 36

Rhynchosia americana, 92; minima,
92

Ribes, 286; amarum, 287; canthari-
forme, 164; cereum, 287, pl. fac-
ing 288; in California, Notes on
the genus, 286; Menziesii, 289,
var. ixoderme, 288, pl. facing 288,
var. leptosmum, 288; petiolare,
289; Roezlii, 289, pl. facing 288,
290; sanguineum, 166, 267

Riccia, 85; fluitans, 242

Richardia braziliensis, 97

Rigiopappus, 73; leptocladus, 73, 74,
(Hee

Rio Grande Valley, Lower, Drainage
of, fig. 45; Geologic map of, fig.
43; Texas, Vegetational survey
of, 41, 77; Vegetation map of,
fig. 47

Rivina humilis, 58, 62, 66, 82, 90

Rock Creek Lake Basin, Inyo County,
California, plants of, a check list,
notice of publication, 239

Romneya, 166

Rollins, R. C., The genus Arabis L.
in the Pacific Northwest, review
of, 37; Glaucocarpum, a new
genus in the Cruciferae, 232

Roripa hispida var. glabrata,
Nasturtium, 82, 91

Rosa, 134; nutkana, 268; gymnocarpa,
268; rugosa, 35

Lomatium,

267;

1938 ]

Rubus nivalis, 268; trivialis, 82, 91

Rudbeckia bicolor, 99

Ruellia californica, 111; intermedia,
97; noctiflora, 97; tuberosa, 52,
97

Rumex Berlandieri, 82, 89; crispus, 89,
266; mexicanus, 60, 89; spiralis,
89

Rumex, The North American species
of, notice of publication, 136

Runyonia longiflora, 64, 89

Sabal Deeringiana, 80; mexicana, 82;
Palmetto, 80; texana, 52, 80, pl.
81, 82, 88

Sabbattia campestris, 95; carnosa, 78,
S01 955.

Sacahuista, 49

Sacahuistal, 49, 54, 77, 80

Sagittaria variabilis, 60, 86

St. John, H., Flora of southeastern
Washington and of adjacent
Idaho, review of, 132; A new
Thalictrum from Mount Rainier,
Washington, 114

St. John, H., and Warren, F. A., The
plants of Mount Rainier National
Park, Washington, review of,
167

Salicornia, 78, 80, 84; ambigua, 80,
89; Bigelovii, 90; herbacea, 80,

90
Salix, 36, 133; arctica var. subcordata,
266; canadensis, 266; lasiandra

var. Lyallii, 266; longifolia, 60,
89; nivalis, 266; petrophila var.
caespitosa, 266; Scouleriana, 266;
sitchensis, 266

Salvia, 197; apiana, 165; ballotaeflora,
96; carnosa subsp. Gilmani, 197;
Clevelandii, 165; coccinea, 58, 96;
columbariae, 208; section Echino-
sphace, 197; section Greeneo-
stachys, 197; Greggii, 82, 96; sec-
tion Jepsonia, 197; Palmeri, 165;
section Parishiella, 197; section
Pycnosphace, 197; review of sec-
tion Audibertia, review of, 196

Salvias, The California, A review of
section Audibertia, review of, 196

Samanea Saman, 121, 123; pollen of,
pl. 125

Sambucus, 261; glauca, 263

Samodia ebracteata, 78, 95

Samolus cuneatus, 95; floribundus, 95

Sanicula septentrionalis, 268

San Diego County, California, Notes
on the flora of, 33; Notes on some
endemics of, 163

Santa Barbara, Battarea phalloides
(Dicks.) Pers., in, 237

INDEX 305

Santa Monica Mountains, California,
Moisture relations in the chapar-
ral of, notice of publication, 101

Sapindus Drummondii, 60, pl. 61, 93

Sarcodes, 1, 7, 8, 9, 10, 11, 140, 146,
150, 153

Sargassum, 71

Saxifraga californica, 33;
folia, 267; rivularis, 267

Schaefferia cuneifolia, 52, 53, 56, 62,
64, 93

Schinus molle, 261

Schrankia, 122; unciniata, 122

Schweinitzia, 138

Scirpus acutus, 242; americanus, 88;
californicus, 88; Hallii, 58, pl. 59,
88; microcarpus, 266; validus, 60,
88

Scutellaria Drummondii, 96;
flora, 269

Sebastiana bilocularis, 111

Sedum integrifolium, 267; variegatum,
163

Seedlings from polyembryonic seeds
of Eugenia Hookeri, 115

Selaginella Wallacei, 264

Selenia, 91

Selloa glutinosa, 99

Senecio, 262; ampullaceus, 99; Ander-
sonii, 239; Elmeri, 270; glabellus,
80, 82, 99; pauciflorus var. fallax,
270; vulgaris, 270

Sequoia gigantea, 1, 285

Serjania brachycarpa, 93; incisa, 52

Sesuvium verrucosum, 78, 90

Setaria geniculata, 87; macrostachya,
58, 64, 80, 83, 87; Scheelei, 87;
setosa, 87

Setchell, W. <A., Nathaniel Lyon
Gardner, 126; A preliminary re-
port on the algae [Templeton
Crocker expedition], notice of
publication, 101

Setchell, W. A. and Gardner, N. L.,
Iridophycus, with special refer-
ence to the South American spe-
cies, notice of publication, 101

Sharsmith, C. W., On the identity of
Claytonia nevadensis Watson, 171

Sharsmith, H. K., The native Cali-
fornian species of the genus
Coreopsis L., 209

Shasta County, Picea Breweriana in,
176

Sherff, EK. E., The genus Bidens, notice
of publication, 136

Shreve, F., The vegetation of the Cape
Region of Baja California, 105

Sida diffusa, 94; filipes, 62, 64, 94;
hastata, 94; paniculata, 58, 94;
spinosa, 94

oppositi-

lateri-

306

Sidalcea spicata, 4

Siderocarpos flexicaulis, 54, pl. 55, 58,
60, pl. 61, 83, 92

Sierra gooseberry, 289

Silene gallica, 262, 263;
267

Silver-tip fir, 2

Silver wattle, 179

Simmondsia californica, 113

Simsia calva, 66, 99

Siphonoglossa dipteracantha, 97

Sisymbrium, 232, 234; altissimum, 267 ;
officinale, 267

Sisyrinchium amoenum, 89; furcatum,
89; longipedunculatum, 78, 89

Sitreavsia pulcherrima, 88

Smelowskia ovalis, 267

Smilacina stellata, 33

Smilax renifolia, 82, 88

Smith, C. P., Species
review of, 236

Solanum carolinense, 97; elaeagni-
folium, 83, 97; Hindsianum, 113;
nigrum, 82, 97, 263; rostratum,
36, 97; triquetrum, 80, 83, 97

Solidago lepida, 270

Sonchus oleraceus, 99, 261, 263, 270

Sophora secundiflora, 92; tomentosa,
80, 92

Sparganium multipedunculatum, 265

Spartina Spartinae, 49, 58, 77, 78, 80,
87

Species Lupinorum, review of, 236

Spergula arvensis, 267

Spergularia rubra, 267

Spiraea Douglasii, 268;
roseata, 268

‘Spirodella, 241; polyrhiza, 242

Sporobolus argutus, 66, 80, 87; Buck-
leyi, 82, 87; cryptandrus, 87;
Wrightii, 66, 87

Spruce, weeping, 176

Spumula quadrifida, 274

Stachys, 270; agraria, 96; bullata,
271, 272; ciliata, 269, 272; Drum-
mondii, 96; Emersoni, 271, 272;
mexicana, 272; palustris, 271;
pilosa, 271; Riederi, 272; rigida,
270, subsp. lanata, 270, 271, 272,
Nutt., Notes on, 270, subsp. quer-
cetorum, 271, 272, subsp. rivularis,
271, subsp. typica, 271, 272

Stanleya, 234

Stebbins, G. L., Jr., An anomalous
new species of Lapsana from
China, 154; Review: Tischler, Die
Bedeutung der Polyploidie fiir die
Verbreitung der Angiospermen,
erliutert an den Arten Schleswig-
Holsteins, mit Ausblicken auf

noctiflora,

Lupinorum,

lucida, 268;

MADRONO

[Vol. 4

The
species of

andere Florengebiets, 198;
western American
Paeonia, 252

Stebbins, G. L., Jr., and Babcock,
E. B., The genus Youngia, notice
of publication, 136

Stellaria media, 262, 263, 267

Stereophyllum Wrightii, 86

Stillingia Torreyana, 93

Stockwell, P., and Wiggins, I. L., The
maritime Franseria of the Pacific
Coast, 119

Streptanthus, 204; batrachopus, 204,
205, 206, pl. 207; Breweri, 205,
206, pl. 207; callistus, 205, 206,
pl. 207, 208; comparative draw-
ings of five species, pl. 207; glan-
dulosus var. pulchellus, 205; his-
pidus, 206, pl. 207, 208; insignis,
206, pl. 207, 208; Nutt., Studies
in the genus, I. Two new species
in the section Euclisia Nutt., 204

Strombocarpa cinerescens, 54, 78, 80,
92

Structure of Allotropa, 137

Stryphnodendron floribundum, 186

Studies in Penstemon VI. The sec-
tion Aurator, notice of publica-
tion, 239

Studies in the genus Streptanthus
Nutt. I. Two new species in the
section Euclisia Nutt., 204

Suaeda, 78, 80, 84; conferta, 78; line-
aris, 80

Subularia aquatica, 267

Suksdorfia ranunculifolia, 267

Swertia perennis, 36

Symphoricarpos albus, 269

Synthlipsis Berlandieri, 91

Systematic study of the genus Erio-
phyllum, notice of publication, 101

Tahoe National Forest, A cranberry
from, 201

Talinopsis frutescens, 64, 91

Talinum, 68; angustissimum, 62, 91;
pulchellum, 68, 69, in Texas, Oc-
currence of, 67; reflexum, 91;
Youngae, 68, 69

Taraxacum laevigatum, 37; officinale,
37, 270

Taxodium distichum,
natum, 62, 86

Tecoma stans, 110, 111

Tephrosia Lindheimeri, 92

Texas, Occurrence of Talinum pul-
chellum in, 67; Vegetational sur-
vey of Lower Rio Grande Valley,
41, 77

Thalictrum, A new, from Mount Rai-

170; + mucro-

1938]

nier, Washington, 114; occiden-
tale, 267; Yrainierense, 114, fig.
114, 115; stipitatum, 115

Thamnosma texana, 93

Thelocactus bicolor, 64, 91

Thelypodiopsis, 232, 234

Thelypodium, 232, 234; suffrutescens,
233

Thlaspi arvense, 33

Thurston, Carl,
Southern California,
publication, 103

Thuya orientalis, 263

Tiarella laciniata, 267

Tillandsia Baileyi, 60, 88; recurvata,
60, 82, 88; usneoides, 60, 82, 88

Tischler, G., Die Bedeutung der Poly-
ploidie fiir die Verbreitung der
Angiospermen, erliutert an den
Arten Schleswig-Holsteins, mit
Ausblicken auf andere Florenge-
biets, review of, 198

Torreya californica, 166, fig. 166, 167

Tournefortia volubilis, 96

Tracyina, 74; a new genus of Aster-
aceae from Northern California,
73; rostrata, 75, fig. 76

Tragia nepetaefolia, 82, 93; urticae-
folia, 93

Tribulus terrestris, 93

Trichachne californica, 87; saccharata,
87

Trichloris mendocina, 66, 87; pluri-
flora, 58, 87

Trichocoronis Wrightii, 99

Tricholaena rosea, 87

Trichoneura elegans, 87

Trichostomum brachydontium, 85

Trifolium dubium, 268; hybridum,
268; pratense, 268; repens, 268

Triodia albescens, 87; mutica, 64, 87;
pilosa, 66, 87

Trisetum cernuum, 265

Trixis californica, 113; radialis, 99

Tiibatulabal ethnography, review of,
236

Tubiflora acuminata, 97

Tuckermannia maritima, 225

Turnera aphrodisiaca, 94; diffusa, 111

Two new species of Linanthus from
western North America, 157

Typha angustifolia, 242; latifolia, 49,
58, 60, 86, 242, 265

Wild flowers of
notice of

Uinta Basin of Utah and Colorado,
Botanical studies in, review of,
196

Ulmus, 262; crassifolia, 60, 82, 89

Undescribed species of Viola from
Utah, 194

Uniola paniculata, 78, 87

INDEX

307

Unusual ascomycete in the shells of
marine animals, notice of publi-
cation, 101

Urtica chamaedryoides, 60, 62, 89

Utah, An undescribed species of Viola
from, 194

Utah and Colorado, Botanical studies
in Uinta Basin of, review of, 196

Utricularia, 103; subulata, 60, 97

!

Vaccinium, 202; macrocarpon, 201;
macrophyllum, 36; oxycoccos, 202,
var. intermedium, 201, 202

Vachellia Farnesiana, 121, 123

Varilla, 64; texana, 64, 84, 99

Vascular plants in Washington, New
records of, 34

Vaseyochloa multinervosa, 80, 87

Vegetation of the Cape Region of
Baja California, 105

Vegetational survey of the Lower Rio
Grande Valley, Texas, 41, 77

Venegasia carpesioides, 33

Veratrum album, 4

Verbascum Thapsus, 269

Verbena, 82, 96; bipinnatifida, 96; cili-
ata, 96; Halei, 80, 83, 96; offici-
nalis, 96; prostrata, 263; xutha, 96

Verbesina encelioides, 53, 64, 66, 83,
99; virginica, 82, 99

Veronica arvensis, 269; Chamaedrys,
36; scutellata, 269

Viae felicitatis: the beginning years
of the California Botanical Soci-
ety, 276

Vicia americana, 268; texana, 92

Viguiera deltoidea, 110, 111; steno-
loba, 54, 56, 58, 99; tomentosa,
110, 111

Vincetoxicum brevicoronatum, 95;
reticulatum, 95

Viola, An undescribed species of, from
Utah, 194; arizonica, 195; Clau-
seniana, 194, 195; lanceolata, 35,
36; mnephrophylla, 195; section
Nomimium, 195; pallens, 36; pa-
lustris, 36; subsection Plagio-
stigma, 195

Voegelin, E. W., Tiibatulabal ethnog-
raphy, review of, 236

Waltheria americana, 94

Warren, F. A., and St. John, H., The
plants of Mt. Rainier National
Park, Washington, review of, 167

Washington, A new Thalictrum from
Mount Rainier, 114; Additions to
our knowledge of the flora of
Mount Baker, 263; Flora of
southeastern, and adjacent Idaho,
review of, 132; New records of

308

vascular plants in, 34; Olympic
Peninsula, A botanical survey of,
review of, 39; Plants of Mount
Rainier National Park, review of,
167

Washingtonia filifera, 262

Wattle, silver, 179

Wedeliella incarnata, 90

Weeping spruce, 176

Weisia Andrewsii, 85

Western American species of Paeonia,
252

Western juniper, 21, 22, 23, 26, Obser-
vations on, 21

Western yellow pine, 22, 26, 27, 169

Wheeler, L. C., Eremocarpus Ben-
tham: Preoccupied? 272

Wiggins, I. L., A resurrection and re-
vision of the genus’ Iliamna
Greene, notice of publication, 102;
Hanging gardens of the Canary
Island date palm, 260

Wiggins, I. L., and Stockwell, P., The
maritime Franseria of the Pacific
Coast, 119

Wilcoxia Poselgeri, 64, 91

Wild flowers of Southern California,
notice of publication, 103

Wilson, A., The California nutmeg
tree in cultivation, 166

Wissadula amplissima, 82, 94

Wolffia, 241, 250; columbiana, 88;
lingulata, 245, 246, 247

Wolffiella, 241; floridana, 241; lingu-

MADRONO

[Vol. 4

lata (Hegelm.) Hegelm., The
flowering of, 241, pl. facing 244,
245, pl. 249, pl. 251, var. minor,
247; oblonga, 241, 244, 245, 246,
247, 248, 250, pl. 251

Woodsia-scopulina, 264

Wynd, F. L., The flora of Crater Lake
National Park, review of, 167

Xanthium speciosum, 99

Xanthoxylum insulare, 82, 93; Pterota,
58, 82, 93

Xylosma celastrinum, 94

Yates, Harry Stanley, 187

Youngia, 156, 157; The genus, notice
of publication, 136

Yucca, 64, 66, 80, 88, 111; Herri-
maniae, 233; rupicola, 88; tenui-
styla, pl. 51, 53, 88; Treculeana,
78, 80, 88

Zacatal, 49, 83

Zacate, 49

Zamia, 277; integrifolia, 170

Zapotal, 66

Zexmenia brevifolia, 99; hispida, 66, 99

Zinnia pumila, 64, 99

Zion National Park, Plants of, review
of, 198

Zizyphus obtusifolia, 53, 54, 56, 58, 62,
64, 66, 80, 82, 83, 94

Zornia bracteata, 96

Zosterella dubia, 86

ERRATA

Page 52, line 19: for tuberculosa read tuberosa.

Page 78, line 10: for D. multiflora read Dondia multiflora.

Page 80, line 9: for leptocephala read leptosepala; line 13, for Hall read
Halei; line 20, for confertifolia read confertiflora.

Page 94, lines 9 and 10: for Amelopsis read Ampelopsis; line 34, for

verticillaria read verticillata.

Page 111, line 41: for Hypis read Hyptis.

Page 126, line 2: for Professor read Associate Professor.

Cover, April 1938, line 3 of Contents: for Caytonia read Claytonia.
Volume 3, page 325, line 44: for 6000 read 3400.

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